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> BULLETIN 


OF 


ENTOMOLOGICAL RESEARCH 


ISSUED BY THE IMPERIAL 
BUREAU OF ENTOMOLOGY. 


EDITOR: THE DIRECTOR. 


VOL XI, 


LONDON: 
Tue Imppria, Bureau or Entomo.noey, 
41, QubEN’s Gate, 8.W.7. 
1920-1921. 


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IMPERIAL BUREAU OF ENTOMOLOGY. 


Honorary Committee of Management. 


VISCOUNT HARCOURT, Chasrman. 


Lieutenant-Colonel A. W. Axooox, C.I.E., F.R.S., Assistant Director, 
Tropical Diseases Bureau. 

Major EK. E. Austen, D.8.0., Entomological Department, British 
Museum (Natural History). 

Dr. A. G. BacsHawe, C.M.G., Director, Tropical Diseases Bureau. 

Major-General Sir J. Rosz Braprorp, K.C.M.G., F.R.S., Royal Society. 

Major-General Sir Davip Bruoz, K.C.B., F.R.S., Chairman of the 
Governing Body, Lister Institute. 

Mr. J. C. F. Frysr, Entomologist to the Ministry of Agriculture. 

Sir Sripnsy F. Harmer, K.B.E., F.R.S., Director, British Museum 
(Natural History). 

Professor H. Maxwett Lerroy, Imperial College of Science and 
Technology. 

The Hon. Sir KE. Luoas, Agent-General for South Australia. 

Dr. R. Stewart MacDoveatt, Lecturer on Agricultural Entomology, 
Edinburgh University. 

Sir Joun McFapyuan, Principal, Royal Veterinary College, Camden 
Town. 

Sir Patrick Manson, G.C.M.G., F.R.S., Late Medical Adviser to the 
Colonial Office. 

Sir Danrzt Morris, K.C.M.G., Late Adviser to the Colonial Office in 
Tropical Agriculture. 

Professor R. Nuwstreap, F.R.S., Dutton Memorial Professor of 
Medical Entomology, Liverpool University. 

Professor G. H. F. Nurraut, F.R.S., Quick Professor of Protozoology, 
Cambridge. 

Professor E. B. Poutton, F.R.S., Hope Professor of Zoology, Oxford. 

Lieutenant-Colonel Sir Davip Pratn, C.M.G., C.I.E., F.R.8., Director, 
Royal Botanic Gardens, Kew. 

Sir H. J. Reap, K.C.M.G., C.B., Colonial Office. 

The Honourable N. C. Rotusouiip. 

Dr. Huan §Scort, Curator in Entomology, Museum of Zoology, 
Cambridge. 

Sir ArtHur EK. Surpruy, G.B.E., F.R.S., Master of Christ’s College, 
Cambridge. 

Mr. R. A. C. Speruina, C.M.G., Foreign Office. 

Sir Stewart Stockman, Chief Veterinary Officer, Ministry of Agriculture. 

Mr. F. V. Turosatp, South Kastern Agricultural College, Wye. 

Mr. C. Warsurton, Zoologist to the Royal Agricultural Society of 

England. 
The Chief Entomologist in each of the Self-governing Dominions 
is ex officio a member of the Committee. 


General Secretary. 
Mr. A. 0. C. Parxtnson (Colonial Office). 


Director and Lditor. 
Dr. Guy A. K. Manrsuatt, C.M.G. 


Assistant Director. 
Dr. 8. A. Neave. 


Head Office—British Museum (Natural History), Cromwell Road, 
London, 8.W.7. 
Publication Office.—41, Queean’s Gate, London, 8.W.7. 


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COME, US. 


ORIGINAL ARTICLES. 


Arrow, GILBERT J. PAGE. 
Some New West Indian Species of the Melolonthid Genus Lachnosterna 189 


AustTEN, Masor E. E. 
The Percy Sladen Trust rape to the Indian Ocean in 1905, and 


1907-1900, © Diptera > Tabanidae?..... es eae 43 
Notes on the Nomenclature of certain African Tabanidae (Sub-family 
Pangoniinae) with Descriptions of a new Genus and new Species.... 139 


BARRAUD, Capt. P. J. 
Mosquitos collected in Palestine and adjacent Territories............ 387 


Bezzi, Pror. M. 
Further Notes on the Lonchaeidae (Dipt.) with Descriptions of New 


Rarer ret APTI SA BIS css. cick ace e oe ving oye pee a cs 199 
Brain, Dr. Cuas. K. ‘ 
ine Woceidac of South Atri¢a Vi. 2.55 dee eee ae then bie 1 


Buxton, P. A. 
A Liparid Moth (Ocnerogyia amanda, Staud.), destructive to Figs in 


Ble aGU OfatA ay 58. onl) soe euetics op on LW Mes Ade, s, Sole NRE 181 
Insect Pests of Dates and the Date Palm in Mesopotamia and else- 
er ere eee may fee 9 sl een het, «age ap 287 


CorBETT, G. H. 
Observations on Cotton Thrips in the Gezira, Blue Nile Province, 
ant) mi See ero cad deo eed Me A pean! fees ah 95 
D EMMEREZ DE CHARMOY, D. 
Notes on Insects accidentally introduced into the Island of Mauritius 171 


D’ EMMEREZ DE Cuarmoy, D. & Tempany, Dr. H. A. 
The Campaign against Phytalus smithi in the Colony of Mauritius.... 159 


DauziEL, Dr. J. M. 


Crab- holes, Trees and other Mosquito Sources in Lagos............. 247 
Epwarps, F. W. 
Notes on the Mosquitos of Madagascar, Mauritius and Réunion...... 133 
On the British Species of Simulium—II. The Early Stages ; with 
Corrections and Additions to PartI.................ceeeee eee: 211 


Fraser, Mayor A. D. 


Notes on the Blood-sucking Flies in North Russia during the Summer 
Se AOLS,..1 hall eet dh. ORRIN cage ns x9 oot dubisinieses 195 


IneraM, Dr. A. & Macrin, Dr. J. W. 8. 


The Karly Stages of West African Mosquitos—V. Culex decens, 
Thee., and Culex imvidioaua, Theo... .... 6.66 e eee ese eee: 105 


(741) Wt. P2/154, 1,000. 7.21. B.&F.Ltd. Gli/i4. 


Vi. CONTENTS 


JACK, RupPErRT W. 
Some Notes and Remarks on the Bionomics of Glossina morsitans.... 113 


LAMBoRN, Dr. W. A. 
Some further Notes on the Tsetse-Flies of Nyasaland.............. 101 
The Habits of a Dipteron predaceous on Mosquitos in Nyasaland .. 279 


Luoyp, Capt. LI. 


On the Reasons for the Variation in the Effects of Formaldehyde as 
Sl OlMGmst OPEL ONISP ted: i naicts «cede ia Peis jet a hd nis. > ee’ <a ee 47 


Macrre, Dr. J. W. S. & Ineram, Dr. A. 
The Early Stages of West African Mosquitos. V. Culex decens, 


Theo., and Culen anerdiosus. Theo. 2 hig pte ob hace ia wicca 105 
MacGrecor, Matcoutm E: 
A New Type of Entomological Killing Bottle...................... 283 
Marsnal., Dr. G. A. K. 
Some New Injurious Weevils. < .0i00 225 sons gids. he PE ee 271 


NEWSTEAD, Pror. R. 
On ‘the Genus Phichotomus. Part VV... os. 32803 eee eee eee 305 


SWELLENGREBEL, Dr. N. H. & SWELLENGREBEL DE GRAAF, J. M. H. 


List of the Anophelines of the Malay Archipelago with special refer- 
ence to Adults and Larvae of new or incompletely described Species 
OP. WAMIGHIES Le efay ooo ste eee cing Were he er ene: ee 2 ae 77 


SWYNNERTON, C. F. M. 


An Examination of the Tsetse Problem in North Mossurise, Portuguese 
Piaet; MMIC a he s2 ow we ee Oe sa Bae oe Beane ee 315 


Tempany, Dr. H. A. & D’ EMMEREZ DE CHARMoy, D. 
The Campaign against Phytalus smithi in the Colony of Mauritius.... 159 


THEOBALD, F. V. 


pnadae (Ob (Ree: soso hes ial de Sake aye, oa ule pic sca SR Ret a 153 

Aiacan' Mehidse ee ary ey pete eek Pee ee aici e sale ee 65 
Uvarov, B. P. 

A preliminary Revision of the Genus Dociostaurus, Fieb. .......... 397 
WituraMs, C. B. 

A Pannier Hopperdover i... eka ees oss hl ee 179 
KETEK, JAMES. 

Anopheles breeding among Water Lettuce. A new Habitat........ 73 

MISCELLANEOUS. 


Collections ‘Tereived..:.. os. .c choses sss nc. ca eee eee 93, 187, 313, 409 


PLATES. 


PAGE. 

lV, South Afticat Goccidae. 0k. 6. he wae cnc tceace facing 42 
V. Views showing captures and destruction of Phytalus smitha 

ERI Ret cas baie gale aces eet a. ke 

VI. A pannier hopper-dozer for catching insects on sugar-cane ,, 186 

wit. opera CUPCUUONIGAG «be ce us leaden cee 5 mie 

VIII. Dates showing ravages of larvae of a Gelechtid Moth ...... i Oe 
IX.—XVII. Types of forest and breeding places of Glossina in 

meromucde Wash ATION, si ool a oie. oy eek ce lads tae dein BOO 

PAGE. 

Charts of Sipane Vlei, 8. Rhodesia...... 2S Sian oa PN cee ar ter 125, 127 


Map illustrating report on the tsetse-fly problem im Portu- 
ene EM, «Sie or 2S Cd ee aay ew Cowes aaa wae <- facing 386 


ERRATA. 


——“-—-— 


Page 159, line 16 for (iii, pp. 93-102) 


o” 


” 


eR id 


9” 


212, 
237, 
254, 
380, 


> 


oT] 


9? 


99 


25 
36 
10 
24 


» writer 
99 D. 


read (viii, pp. 93-102) 


9? 


3° 


» Rev. App. Ent. i 


». (PL 


, fig. 46) 


winter 

Dr. | 

Rev. App. Ent., B, i 
(Pl. XVII, fig. 1) 


- 


VOL. XI. Part 1.—pp. 1-94. 
; AUGUST, 1920, 


BULLETIN OF — 
ENTOMOLOGICAL 
RESEARCH 


rss 
sU7) tee 
SMe. . ) 
L 2 « wf AW*ga 4 . 
APRi¢ 1923 


\ . 


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hiya Qf 
Luo; 


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NiAN OS 


LONDON. 
THE IMPERIAL BUREAU OF ENTOMOLOGY, 
88, QUEEN’S GATE, S,W. 7. 


Price 5s. net. 


All Rights Reserved. 


IMPERIAL BUREAU OF ENTOMOLOGY. 
HON ORARY COMMITTEE OF MANAGEMENT 


‘THE VISCOUNT HARCOURT, Cbafrman. 

Lrevt.-Cotonen A. ALCOCK, C.LE., ERS. 

Mayor E. E. AUSTEN, D.S.0. 7 a ff 

Dr. A. G. BAGSHAWE, C.M.G: | 

Major-GEenerat Siz JOHN R. BRADFORD, K.C.M.G., E.B.S. 
-  Masor-GenERaL Siz DAVID BRUCE, K.C.B., F-.R.S. 

Mr; J: C. F. FRYER. | 

Siz SF. HARMER, K.B.E., FBS. | 

Pros. H. MAXWELL LEFROY. i 

Hon. E. LUCAS. 

“Dr. R: STEWART cachoseate 

Sir JOHN McFADYHAN. : 

Str PATRICK MANSON, G.C.M.G., F.R.S. | 

Siz DANIEL. MORRIS, K.CMG ee 

Pror. R. NEWSTEAD, F.R.S. : 

Pror. G. H. F.. NUTTALL, F-.R.S. 

Pror. E. B. POULTON, FBS. 
Lizut,-Cotonet Str DAVID PRAIN, C.M. G., Ci LE., E.R.S. 

Sir H. J. READ, K.CMG., CB. 4 

Tun Hon. N. C. ROTHSCHILD. : | 

De. HUGH SCOTT. — | : * 

Sin A. E, SHIPLEY, G.B.E., PRS. | | ) 

Mr. R. A. C. SPERLING, C.MLG. om 

Sm STEWART er 

Mr. F. V. THEOBALD. 

Mr. C, WARBURTON, | 


| Director. 
| Dr. GUY A. K. MARSHALL, C.M.G. 


Fssistant Director. ie 
Dr. S& A. NEAVE, ~ | 
Secretary. | 
Mr. A. C. C. PARKINSON. 


IMPERIAL BUREAU OF ENTOMOLOGY. 


BULLETIN 


OF 


mNTOMOLOGICAL RESEARCH. 


Vou. XI. 


1920. 


THE COCCIDAE OF SOUTH AFRICA—V.* 
By Cuas. K. Brain, D.Sc. 
Division of Entomology, Pretoria, South Africa. 


Subfamily LecanimnaE 
Key to Genera 
Genus Lecanium, Burm. 


Saissetia, Deplan. 
Hemilecannum, Newst. 


Aclerda, Sign. 


Allopulvinaria, gen. nov. .. 


Protopulvinaria, Ckll. 
Pulvinaria, Targ. 
Ceronema, Mask. 
Tnchtensva, Sign. 
Filippa, Targ. 
Conofilippra, gen. nov. 
Ceroplastes, Gray 
Inglisia, Mask. 
Cryptinglisia, Ckll. .. 
Parafairmairea, Ckll. 
Ceroplastodes, Ckil. .. 


Idiosatssetia, gen. nov. 


Membranaria, gen. nov. 


(Piatses I-LV.) 


CONTENTS. PAGE 


25 
26 
36 
38 
39 
40 
40 
4] 


Subfamily LECANIINAE. 

This subfamily comprises a large number of genera, which are, as a rule, easily 
distinguished by the presence or absence of secretionary matter in one form or 
another. All agree, however, in having the anal ring with a number of hairs and, 
further, in having the hind margin cleft. The anal ring is covered above by a pair 
of more or less triangular plates which are usually densely chitinised. The eighteen 
genera known to be represented in South Africa are tabulated below :— 

*For Part II, sce Bull. Ent. Res. ix, p. 107; Part III, op. cit. ix, p. 197; Part IV, 


op. cil. X, p. 95. 
(681) Wt.P4/140. 1,000 7.20 B.&F.,Ltd. Gp. 11. A 


2 CHAS. K. BRAIN. 


Key to South African Genera. 
A. Female naked, or only covered by a very thin, inconspicuous film of secretion. 
B. Adult 9° with well-developed legs and antennae. 
C. Adult 2 with posterior extremity cleft. 
(1) Female flat or slightly convex, derm not very hard when mature, without 


polygonal areas containing pits .. A Lecanum, Burm., p. 3. 

(2) Female becoming hard and convex at maturity, derm with + oval or 
polygonal areas containing _ pits ae Sarssetra, Deplan., p. 2. 

CC. Adult © with posterior Baas not cleft; female with four large 
groups of spinnerets .. ; a .. Hemilecanmum, Newst., p. 14. 
BB. Adult 92 without legs or ae - a .. Aclerda, Sign., p. 15. 


AA. Female secreting a greater or less amount of cottony or powdery matter. 
B. Female naked above, secretion beneath or behind the insect. 
C. Female with only a narrow fringe of secretion. 
(1) Female very convex, dark red, abdominal segmentation very distinct 
Allopuluinaria, gen. nov., p. 16. 
(2) Female flat or slightly convex, segmentation not pronounced, 
Protopulvinaria, CkIL., p. 16. 
CC. Female secreting a definite protruding ovisac Pulvinaria, Targ., p. 17. 
BB. Female partly covered with secretion. 
(1) Adult 2 covered above, except for the median area, with a thick coat of 
wavy threads which project beyond the margin in all directions 
Ceronema, Mask., p. 22. 
(2) Adult 92 enclosed in a felted sac, ne the cephalic extremity which 
is + exposed Bs ; . .Inchtensia, Sign., p. 22. 
BBB. Adult 9 entirely enclosed. 
C. Sac fitting closely to body of adult 9 at maturity and later serving as 
ovisac ; insect leaf-infesting .. 2 . Filipyra, Targ., p. 23. 
CC. Sac felted, stout, conical, insect ah ee Ganctienal gen. nov., p. 25. 
AAA. Female secretion waxy, glassy or horny. 
B. Covering of 2 consisting of wax, generally soft and thick ; no marginal fringe 
nor marginal processes ; female with a + conspicuous caudal process visible 
on removing the wax .. ie Ceroplastes, Gray, p. 26. 
BB. Covering of 9 glassy, or at ie pide, thin. 
(1) Scale divided into Se and striated with rows of air-cells ; not causing 
galls vas : me Inglisia, Mask., Dp. 36. 
(2) As in Inglisra iit eateaae aaille on ene of host-plant 
Cryptinghsta, Ckll., p. 38. 
(3) Scale divided into two halves not striated with air-cells, but with grooves 
radiating from apex bi ne <p Parafairmairea, Ckll., p. 39. 
(4) Scale not divided into plates nor two halves, but rough and beset with 
protuberances, legs and antennae well- developed, anal cleft normal 
Ceroplastodes, Ckll., p. 40. 
(5) Scale not divided into plates nor two halves, + smooth, legs and antennae 
rudimentary, anal cleft lateral .. al diosaissetia, gen. nov., p. 40. 
(6) Similar to Pulvinaria but ovisac consisting of horny membrane instead 
of cottony matter .. ye ips .. Membranama, gen. nov., p. 41. 


THE COCCIDAE OF SOUTH AFRICA. S 


Genus Lecanium, Burm. 


Adult 9 never hemispherical nor highly convex, derm remaining comparatively 
soft, without polygonal areas; usually light in colour and oval; legs and antennae 
present, well-developed. 


191. Lecanium hesperidum, Linn. 
Coccus hesperidum, Linn., Syst. Nat. Ed. x. 1, p. 455, 1758. 
Chermes hesperidum, Geoff., Hist. Abr. Ins. i, p. 505, 1762. 
Calypticus hesperidum, Costa, Faun. Reg. Nap. Cocc., p. 8, 1835. 
Calypticus laevis, Costa, Faun. Reg. Nap. Cocce., p. 8, 1835. 
Calymnatus hesperidum, Costa, Nuov. Osserv., p. 22, 1835. 
Coccus patelliformis, Curt., Gard. Chron., p. 517, 1843. 
Chermes lauri, Bdv., Ent. Hort., p. 340, 1867. 
Lecanium platyceru, Pack., Rep. Mass. Bd. Agr., p. 260, 1870. 
Lecanium angustatum, Sign., Ann. Soc. Ent. Fr. (5) i, p. 398, 1873. 
Lecanvum maculatum, Sign., Ann. Soc. Ent. Fr. (5) i, p. 400, 1873. 
Lecanium alienum, Dougl., Ent. Mon. Mag., xxi, p. 77, 1886. 
Lecanium depressum var. simulans, iene Ent. Mon. Mag. xxiv, p. 28, 1887. 
Lecanium minimum, Newst., Ent. Mon. Mag., xxvu, p. 141, 1892. 
Lecanwum assimile var. amaryllidis, Ckil., Tr. Am. Ent. Soc. xx, p. 53, 1893. 
Lecamum terminaliae, Ckil., Jl. Inst. Jamaica, 1, p. 254, 1893. 
Lecanium nanum, Ckll., Psyche, vii, Suppl., 1, p. 19, 1896. 
Lecanium flaveolum, Ckll., Psyche, viii, pp. 52, 53, 1897. 
Lecanvum minimum var. pinicola, Mask., N.Z. Trans. xxix, p. 310, 1897. 
Lecanwum ventrale, Ehrh., Can. Ent. xxx, p. 245, 1898. 
Lecanvum (Calymnatus) hesperidum pacificum, Kuw., Jl. N.Y. Ent. Soc. x, 
p- 30, 1902. 
Lecanvum signiferum, Green, Coce. of Ceylon, pt. ii, p. 197, 1904. 


“ Adult female bright yellow or greenish-yellow, minutely speckled with red- 
brown, the specks sometimes agglomerated into transverse bars, especially on the 
median abdominal region ; in other parts tending to form dotted lines radiating 
from centre to margin. In older examples the ground-colour may be ochreous or 
pale fulvous ; and the maculation may form a broad median fascia. Under surface 
of older examples with a deep purple-brown or red patch covering the median 
abdominal area, becoming concave and forming a shelter for the young larvae. 
Dried specimens straw-coloured and much wrinkled. Form oblong-oval, often 
very irregular in outline ; narrowest in front ; more or less concave above according 
to age. In some individuals, generally on those protected by some shelter, I have 
noticed a double median longitudinal series of raised glassy points; but they 
appear to be very brittle and easily lost.’ (Green). 

Length, 25-5 mm.; breadth, 15-3 mm. 

The antennae are 7-jointed, with the 3rd, 4th and 7th joints subequal. The 
tange of measurements from South African material is as follows :—(1) 27-35, 
(2) 30-37, (3) 51-61, (4) 54-65, (5) 20-24, (6) 17-27, (7) 44-51. 

The anal plates are about 155y long. 

(681) A2 


4 CHAS. K. BRAIN. 


The hairs on the antennal joints are exceptionally thin and are distributed, 
2 on joint I, 2 on II, 2 on IV, 1 on V and VI, 8 or 9 on VII. 

The marginal spines are short, widely spaced, simple, and nearly always have their 
tips recurved. There are three stigmatic spines, laterals short, median about 33 
times as long as laterals. The stigmatic clefts are but slightly recessed, with a few 
simple glands. 

Habitat: On citrus and a large variety of cultivated plants common throughout 
the Union. 

Collection Nos.: 110 and 136. 


192. Lecanium africanum, Newstead. 


Lecanium viride var. africanum, Newst., Ent. Mon. Mag. (2) ix, p. 95, 1898. 

Lecanium viride (Green), Newst., Bull. Ent. Res. i, p. 187, 1910. 

Lecanium africanum, Newst., Bull. Ent. Res. vu, p. 357, 1917. 

“Female adult. Colour of dried specimens often yellowish green; others are 
bright ochraceous, straw-coloured or pale reddish-brown; eyes black. Dorsum, 
especially in the younger forms, with a series of black markings, often forming a 
narrow loop-like pattern ; these markings are however rarely present in very old 
examples. Form oblong oval, but the outline is often irregular; narrowed in 
front and moderately convex, but the margins broadly flattened, especially the 
cephalic portion. Antennae of eight segments, rarely of seven; formula of the 
former: 3 (2, 4, 5, 8) 1 (6,7); there is a very long hair on the 2nd and 5th segments, 
and several shorter ones on the 8th, 6th and 7th, each with a distinctly stouter hair. 
Legs normal, though the anterior tarsi sometimes exhibit a faint dorsal constriction. 
Anal lobes attenuated distally, inner margin longest; apices with a few fine short 
hairs. Anal cleft slightly less than one-third the length of the body. Stigmatic 
cleft very slight, sometimes scarcely visible; spines three in number, rather small, 
the central one being about three times the length of the laterals. Marginal spines 
small, generally slightly curved and faintly fringed distally. There are three to 
four hairs of varying length near the attachment of each of the antennae, and usually 
four rather longer and stouter ones just in advance of the anal lobes. Derm cells 
oval and markedly distinct in stained preparations, but scarcely visible in unstained 
specimens.”’ (Newstead). 

Habitat: On citrus, Pietermaritzburg, Natal, and Nelspruit, Transvaal. On 
coffee, Natal coast (Fuller). 

Collection No.: 106. 


193. Lecanium ehretiae, sp. n. 


Adult 2 somewhat similar to L. hesperidum but rather less convex and much darker 
in colour, dark brown to blackish, 3°6 mm. long by 1:8 mm. broad, regularly oval. 
A single fringe of very short hairs is noticeable around the margin when examined 
with a hand lens and the surface of the body then appears mottled greyish brown 
in colour; the anal plates are bright brown. The dorsal surface is slightly shiny, 
with very indistinct and indefinite ridges and depressions. . 


THE COCCIDAE OF SOUTH AFRICA. 5 


The antennae are 7 or 8—jointed, most often 7. The following range is obtained 
from a series of eight measurements :—7-jointed : (1) 24-34, (2) 54-61, (3) 54-61, 
(4) 68-75, (5) 24-31, (6) 14-24, (7) 34-41; 8-jointed : (1) 34, (2) 51, (3) 65, (4) 51, 
(5) 30, (6) 20, (7) 21, (8) 41 u. 

Leg I: coxa 68, femur + tfochanter 170, tibia 126, tarsus 75, claw 20y. 
Tarsal digitules very slender, hardly perceptibly clubbed. Trochanter with an 
apical spine about 85y long. 

_ Scattered over the integument, particularly around the antennae and anal plates, 
are numerous hairs, the longest of which may reach 80y in length. 

The marginal spines are comparatively long (80x), slender, slightly curved, not 
truncate. They average approximately 54u apart. The stigmatic cleft has two 
short spurs and a median longer one (40u). The anal plates are comparatively 
short (142) and broad (170y across the outer angles). The anal ring has six long 
hairs. The derm is apparently clear, without glands. 

Habitat : On Ehretia hottentottica, Burch. ; collected by the writer at Brooklyn, 
Pretoria, October 1914. 

Collection No.: 117. 


194. Lecanium pumilum, sp. n. 

Adult 2 small, about 2 mm. long, and 1°5 mm. broad, moderately convex, dark 
reddish brown in colour, distinctly red when younger. 

Antennae 7- or 8-jointed, more often 8. The following represents the range 
obtained from measuring the antennae of five insects :—7-jointed: (1) 20-24, 
(2) 34-87, (3) 51-54, (4) 27, (5) 24, (6) 17-20, (7) 41-54; 8-jointed: (1) 24-27, 
(2) 27-44, (3) 48-51, (4) 17-24, (5) 24-34, (6) 17-20, (7) 17-20, (8) 37-44. 

Leg I: coxa 51, femur + trochanter 153, tibia 95, tarsus 61, claw 171. 

The anal plate is about 150u long. The anal ring is produced on a delicate fluted 
tube ; it bears 8 long bristles and several shorter ones. The margin has the usual 
series of simple spines about 41u long. Stigmatic cleft with 3 straight, moderately 
stout spines; laterals about 24u long, median about 71u. Immediately within the 
“margin are numerous scattered small simple gland pores, many with short sharp 
spines. 

Halntat : On the stem of a native shrub, covered by a carton shelter constructed 
by ants ; collected by C. P. v.d.Merwe at Robertson, C.P., May 1915. 

Collection No.: 125. 


195. Lecanium elongatum, Sign. (Plate i, fig. 233.) 

Lecanium elongatum, Sign., Ann. Soc. Ent. Fr. (5) i, p. 404, 1873. 

Lecanium longulum, Dougl., Ent. Mon. Mag. xxiv, p. 97, 1887. 

Lecanium chirvmoliae, Mask., N. Z. Trans. xxii, p. 137, 1889. 

Lecanium ficus, Mask., Ent. Mon. Mag. xxxiil, p. 243, 1897. 

Lecanium frontale, Green; Cocc. Ceylon, pt. iii, p. 192, 1904. 

“Female dingy pale yellowish-grey ; elongate, narrow, ends broadly rounded, 
side margins slightly curved out, not recurved; surface smooth, transversely 
arched, longitudinally level, semi-cylindric, not carinate, a band of fairly dark 


6 CHAS. K. BRAIN. 


reticulation along the sides, whence, in some examples, faint dark lines radiate to 
the margin ; the disc occupied with a long, pale, clear, oval spot ; or in some mature 
specimens the scale (female) is unicolorous yellow-brown, the dorsal pale spot 
partly or wholly covered, and on the sides minute pale dots in place of reticulation. 
Underside all pale, a broad space all round the insect, a conspicuous blackish eye- 
spot above each antenna.” (Douglas). 

Antennae long, of 8 segments, range in mw as follows :—(1) 34-40, (2) 51-58, (3). 
68-82, (4) 54-58, (5) 41-54, (6) 24-31, (7) 20-31, (8) 34-41. 

Anal plates longer than in hesperrdum (170), with the outer angles more rounded. 
Marginal spines a little longer than in hesperidum (34), widely spaced, averaging 
70-85 apart. Usually simple, but some with apex broadened and slightly branch- 
ed and recurved. Integument, when fully chitinised, with scattered, small, oval, 
clear spaces. Stigmatic spines (3); laterals longer, straighter and more pointed 
than in hesperrdum, median about four times the length of laterals, broad at the 
base and tapering to an acute point. 

Habitat: On Acacia melanorylon; collected by A. Kelly, Joubert Park, 
Johannesburg, May 1915. 

Collection No.: 111. 


196. Lecanium pseudelongatum, sp. n. 

Adult 9 similar to elongatum but slightly less convex and darker in colour. The 
antennae are 8-jointed as in elongatum, but joints 3 and 4 are longer and 5 
shorter, giving the following range :—(1) 34-40, (2) 41-51, (3) 59-68, (4) 44-6], 
(5) 61-71, (6) 24-34, (7) 24-27, (8) 37-41. 

The most striking difference, however, is found in the marginal spines which 
are long (58) and close together, averaging 34u apart. Stigmatic spines (3) 
with the laterals relatively short and slender and the median long, curved and linear. 

Habitat : On native thorn tree, Acacia caffra (?), Pretoria; collected by the writer, 
September 1914. 

Collection No.: 116. 


197. Lecanium filamentosum, N ewst. 

Lecanium (Eulecanium) filamentosum, Newst., Bull. Ent. Res. iv, p. 74, 1913. 

Adult 2 about 6 mm. long and 4 mm. broad, flatly convex with a slight median 
ridge. The colour of dried material is pale brown, with median area white as though 
covered by a dense layer of white secretion, which gradually fades towards the 
margin where numerous black dots are evident. The margin is faintly indentate, 
this character being intensified by the lines of black dots running to the edge and 
the distinct marginal glassy fringe. Seen from below the insect is brown, with four 
conspicuous stigmatic bands. 

The antennae are 8-jointed with joint 3 remarkably long. A series of ten 
measurements gave the following range in wu :—({1) 57-65, (2) 68-82, (3) 150-187, 
(4) 75-88, (5) 65-75, (6) 34-44, (7) 31-37, (8) 54-58. 

The integument is characterised by a few scattered, short, conical spines and a 
number of small, indistinct, “ rosette’ gland openings. The anal plates are about 
220u long each, with a stout apical spine and several others on the disc. 


THE COCCIDAE OF SOUTH AFRICA. 7 

The marginal spines are comparatively short (37), truncate, with the extremity 
slightly forked. They are set at an average distance of about 50u apart. The 
stigmatic clefts are deeply indented but not recessed ; each with about 8 spines, 
of which the laterals are comparatively long and stout (60u); the middle spine 
is only a little longer. but stouter. The tarsal digitules are very long and slender ; 
those of the claw are short and very broad. 

Habitat : On fig, Cathcart, C. P.; collected by L. J. Botha, July 1918. 

Collection No.: 314. 


198. Lecanium proteae, sp. n. 

Adult 9° about 6 mm. long and 4:5 mm. broad (longest specimen seen reaches: 
8 mm. by 6 mm. broad); colour, when alive, asphodel green (Ridgway) with 
very narrow margin of pale dull green yellow. The dorsum is dull coriaceous, giving 
the insect the colour and appearance of the leaf, the green body corresponding with 
the blade and the yellowish margin with the midrib or edge of the leaf. 

The anal plates are small, yellowish in colour, except in old specimens, in which 
they are tinged with brown. 

In form the insect is uniformly oval, except when situated near the midrib or 
edge of a leaf, when the form is irregular. Above, the body is moderately convex. 
Below, the insect is uniform green in colour. In boiling KOH it turns to orange 
yellow then reddish brown. After treatment the derm is colourless. 

Anal plates about 205 long, outer angles rounded, apices attenuated, with two 
apical spines, one subapical and one on the disc at about one-third the length from 
the apex. Anal ring with 6 long hairs. On the dorsum, immediately behind the 
anal opening, are two scattered groups of “rosette”’ glands, and there is a further 
transverse series on the segments immediately anterior to the two groups. The 
integument is otherwise clear, but has scattered hairs of varying lengths. 

Stigmatic clefts shallow, with short, varying, clubbed processes. 

Antennae usually 7-jointed with the fourth showing a pseudarticulation ; some- 
times 8-jointed; 7-jointed form : (1) 27, (2) 37-41, (3) 51-58, (4) 44-48, (5) 20-24, 
(6) 20-24, (7) 41-54; 8-jointed form: (1) 27, (2) 41, (3) 61, (4) 34, (5) 31, (6) 20, 
(7) 20, (8) 51y. 

Leg IL: coxa 54 by 65, femur + trochanter 127 by 37, tibia 95, tarsus 17, claw 20u. 

Marginal spines very short (17) at very wide intervals, averaging at least 80u 
apart, simple, truncate. 

Male scale of the usual type, 2-0 mm. long and 1:2 mm. broad. Head and body, 
without genital sheath, 1-6 mm. long. Head and prothorax rufous to nearly black, 
glistening ; body reddish, tip of abdomen and spine lighter, yellowish ; legs dark, 
slightly darker than antennae. Caudal filaments, two, white, about as long as the 
head and body without antennae. Wings whitish, 1-6 mm. long, mealy, with a 
subcostal reddish line running three-fourths of length of wing from the base. 
Antennae 1 mm. long, of ten joints. 

Larva, newly emerged, 0°5 mm. long and 0°25 mm. broad, orange yellow, very active. 

Habitat : On leaves of Protea, Pretoria ; common. 

Collection No. : 108. 


8 CHAS. K. BRAIN. 


199. Lecanium wistariae, sp. n. 

Adult 9, mounted specimens about 3°4mm. long by 2 mm. broad, elongate, 
tapering to each end. The following description, taken from the Cape Journal, 
was made from fresh material :— 

“Young appearing August 15th; females apparently viviparous, since, while 
bodies are filled with eggs, only living young are seen under the scales. 

“Female long oval, highly convex. Ground-colour yellow obscured by 
reticulations of black; the black enclosing minute, almost round patches and 
bounding larger patches of irregular shape and size; in a few specimens there is 
an obscure median stripe of yellow, but this is never well defined and generally 
altogether lacking. A tinge of red or rose is apparent about the whole margin of 
some specimens and this in a few extends as a flush over the whole dorsum.” 

Antennae 8-segmented; range: (1) 30-34, (2) 36-42, (3) 54-62, (4) 34-44, 
(5) 48-52, (6) 26-30, (7) 20-24, (8) 34-38. 

Leg I: coxa 54 by 70, femur + trochanter 170 by 40, tibia 130, tarsus 68, claw 20u. 
The tarsal digitules are slender and clubbed. 

Margin with a series of slender spines, 44 long, not truncate. Stigmatic cleft 
spines similar but stouter. Anal plate 125, long. 

Habitat: On wistaria, Uitenhage, collected August 1901; Cape No. 1286. 
Thickly clustered on thin twigs of plant ; also said to be on rose and Australian 
myrtle. 

Collection No. : 113. 


200. Lecanium durbanense, sp. n. 

Adult 2 about 5 mm. long and 4 mm. wide, broad egg-shaped, with the extremities 
broadly rounded. The anterior end narrows perceptibly just in front of the middle. 
Body flat, mahogany-brown, with the margins and a faint median ridge somewhat 
darker, glossy. 

When mounted, the most striking character under the low power is the chitinisation 
of the dorsum. In a specimen measuring 5 mm. in length a conspicuous chitinised 
line proceeded from the anal lobes along the middle of the dorsum to the level of the 
mouth-parts. A faint chitinised band was also noticeable around the margin. 

The antennae are 7-or 8-jointed, with the following range in yw :—8-jointed series : 
(1) 27-34, (2) 34-41, (3) 37-51, (4) 34-48, (5) 41-44, (6) 27, (7) 27-31, (8) 44-51; 7- 
Jointed series: (1) 27-34, (2) 37, (3) 37-51, (4) 37-41, (5) 41-44, (6) 37-51, 
(7) 48-51. 

Leg I: coxa 88, femur + trochanter 187, tibia 119, tarsus 82, claw 20u. 

Anal plates about 156 long, comparatively slender. The marginal zone appears 
more chitinous than the median area with the exception of the ridge previously 
mentioned. The chitin of the zone appears stippled ; that of the median ridge has 
numerous small oval transparent spots. The marginal spines are very wide apart 
(110), and comprise a single row of short conical spines. The stigmatic clefts are 
recessed in the form of narrow deep cups ; each contains two short (27,) blunt spurs 
at the base. i 


THE COCCIDAE OF SOUTH AFRICA. 9 


Scattered over the integument are a few comparatively long hairs, which are 
particularly noticeable in the region of the antennae. 

Habitat: On leaves of plant, species undetermined, Durban ; collected by C. P. 
v. d. Merwe, April 1916. 

Collection No.: 126. 


Genus Saissetia, Deplan. 
Adult 2 usually very convex or hemispherical, and the integument very dense, 
dark in colour, and hard at maturity, with cell-like markings ; legs and antennae 
well-developed. 


201. Saissetia hemisphaerica (Targ.). 
Lecanium hemisphaericum, Targ., Studi sul. Coce. pp. 26, 27, etc., 1867. 
Chermes anthuru, Bdv., Ent. Hort. p. 328, 1867. 
Chermes filicum, Bdv., Ent. Hort. p. 325, 1867. 
Chermes hibernaculorum, Bdv., Ent. Hort. p. 337, 1867. 
Lecanium coffeae, Sign., Ann. Soc. Ent. Fr. (5) i, p. 435, 1873. 
Lecanium beaumontiae, Dougl., Ent. Mon. Mag. xxiv, p. 95, 1887. 
Lecanium clypeatum, Dougl., Ent. Mon. Mag. xxv, p. 58, 1888. 
Lecanium hemisphaericum var. hibernaculorum, Ckll., Bull. Bot. Dept. Jamaica, p. 
71, 1894. 

Lecanium hemisphaericum var. filicum, Green, Ent. Mon. Mag. xxxili, pp. 70, 77, 
1897. 

Lecanium (Saissetia) coffeae var. clypeatum, Ckll. & Parr., The Industrialist, p. 164, 
1899. 

Saissetia hemrsphaerica, Ckll., The Ent. Student, u, p. 32, 1901. 

Coccus coffeae, Kirkaldy, Fauna Haw. in, pt. 2, p. 105, 1902. 

Adult 9 approaching hemispherical, ovate, with the margins somewhat flattened ; 
. dorsum smooth, shining, light to red-brown, about 2 to 4 mm. long, 1 to 2°5 mm. 
broad, and 1:5 to 2 mm. high. In the young forms an indistinct H is sometimes 
indicated on the dorsum, but this disappears in the adult, which is thus readily dis- 
tinguished from S. oleae. 

Integument with numerous clear, oval derm cells. Antennae 8-jointed, with the 
following range in w: (1) 37-45, (2) 47-54, (3) 74-81, (4) 46-51, (5) 44-53, (6) 27-34, 
(7) 24-30, (8) 48-54. , 

Leg III: coxa 110, femur + trochanter 255, tibia 185, tarsus 88, claw 241. 

Anal plates about 136 long, triangular, with rounded corners. Marginal spines 
flattened at the ends, which are serrated in a variety of forms, some simple ; stig- 
matic spines all strong and blunt ; median longer than laterals. 

Habitat: On pot-plants, Capetown, Grahamstown (C. P.), Durban and Pieter- 
maritzburg (Natal), and Pretoria and Johannesburg (Transvaal). 

Collection No.: 120. 


10 CHAS. K. BRAIN. 


202. Saissetia oleae (Bernard). 

Chermes oleae, Bern., Mem. d’Hist. Nat. Ac. Marseille, p. 108, 1782. 

Coccus olea, Oliv., Ency. Meth. vi, p. 95, 1791. 

Coccus palmae, Haw., Tr. Ent. Soc. Lond. p. 307, 1812. 

Coccus testudo, Curt., Gard. Chron. p. 444, 1843. 

Coccus cycadis, Bdy., Ent. Hort. p. 323, 1867. 

Lecanium cassiniae, Mask., N. Z. Trans. xxii, p. 15, 1890. 

Lecanium oleae var. testudo, Ckll., Check List, p. 331, 1896. 

Lecanvum oleae var. mirandum, Ckll. & Parr., Biol. Centr. Am. 11, pt. 2, p. 12, 1899. 

Coccus oleae, Kirkaldy, Fauna Haw. i, pt. 2, p. 106, 1902. 

Adult 9 short ovate, high convex to almost hemispherical, 2°5 to 4 mm. long, 1:5: 
to 3mm. broad, and 1:5 to 2°5 mm. high. The dorsum has one long:tudinal and two 
transverse ridges forming a distinct H, dark brown, often dotted with minute flecks. 
of white wax. Derm with elongate cells, each enclosed in an irregular, polygonal 
tessellation. 

Antennae 8-jointed, the segments measuring in w: (1) 34, (2) 44-48, (3) 66-75, 
(4) 35-42, (5) 24-37, (6) 24-32, (7) 24-32, (8) 44-51. 

Leg I: coxa 78, femur + trochanter 170, tarsus 85, claw 24u 

Anal plate about 176u long. Marginal spines simple or flattened at apex. 

Habitat: On a variety of a fairly common throughout the Union. 

Collection Nos.: 122 and 122 

A distinct variety of this mene characterised by being aes and flat, without. 
the dorsal H, is often found associated with typical oleae on oleander at Capetown. 
Its microscopic characters agree with oleae with the exception that antennal 
joints 2 and 4 are a little longer (Coll. No. : 122a). 


203. Saissetia nigra (Nietn.). 

Lecanvum nigrum, Nietn., ‘“ Enemies of Coffee-tree,” p. 9, 1861. 

Lecanium depressum, Targ., Studii sul. Coce., p. 29, 1867. 

Lecanium begoniae, Dougl., Ent. Mon. Mag. xxviii, p. 209, 1892. 

Lecanium nigrum var. depressum, Ckll., Check List, p. 332, 1896. 

Saissetia depressa, King, Psyche, ix, p. 296, 1902. 

Saissetia nigra, King, Psyche, ix, p. 296, 1902. 

Sarssetia nigrella, King, Psyche, ix, p. 296, 1902. 

Coccus mgrum, Kirkaldy, Fauna Haw. ii, pt. 2, p. 106, 1902. 

Adult 9 long oval to broad ovate, low convex, shining black, 3 to 4 mm. long. 
Marginal spines scattered, simple, about 40u long. Antennae of 7 or 8 segments, 
7-jointed series with 4 longest, 8-jointed series with 3 longest; range in u :— 
8-jointed series : (1) 27, (2) 37-41, (3) 58-68, (4) 34-44, (5) 34-41, (6) 24, (7) 24, (8) 
51-54 ; 7-jointed series: (1) 27, (2) 34, (3) 54, (4) 68, (5) 24, (6) 20, (7) 58. 

Leg I: coxa 58, femur + trochanter 170, tibia 116, tarsus 68, claw 20. 

Anal plate about 170u long. 
Habitat: On Ficus spp., Bayville, C. P., and Natal coast. 
Collection No. : 123. 


THE COCCIDAE OF SOUTH AFRICA. lI 


204. Saissetia perseae, sp. n. 

Ova and larvae small, purplish red in colour, found beneath dry body of adult Y 
attached to mid-rib of leaf. Larvae about 0°35 mm. long and 0:18 mm. broad, 
almost oval, slightly broadest between the second and third pairs of legs, broadly 
rounded in front and regularly cleft behind, with two caudal spines equal in length 
to the greatest width of the body. The eyes are deeply pigmented. 


The antennae are six-jointed; segment 2 a little shorter than 1; 3 is long 
(27-30), almost equal to 6; 4 and 5 are shorter and almost equal (13 and 15,). 
There is a distinct notch near the distal end of 3, and another at about the middle 
of 5, from which points long hairs arise. The terminal segment has a long apical 
spine, two others of about three-fourths its length, and several shorter hairs. Claws 
simple; upper digitules long, straight, with small globular knobs ; lower digitules 
shorter, also with small clubs. Margin with thin short spines ; at level of spiracles 
on each side there are two stout blunt spines, about 15 long, pointing obliquely 
backwards. Anal bristles reaching level of hind-margin, or nearly so. 


Adult 9 4:2 mm. long, 3 mm. broad, pointed at each end, but more so in front. 
with the dorsum quite flat. In colour it is blackish brown, and of a dull matt 
appearance, entirely without design or marginal appendages. The margin is 
entire, with a very slight notch at the two spiracles on each side. Seen from below 
the extreme margin is thin; the legs and antennae pale, and there are four faint 
lines, two on each side, representing stigmatic bands. There is no trace of an anal 
cleft when examined with a hand-lens. When crushed the body-contents are 
purple in colour. Placed in hot KOH solution the body becomes hard and curls so 
much that it is difficult to make a satisfactory mount. 

The integument on the dorsum is extremely dense, hard and brittle when cleared ; 
that of the venter is thin, hyaline, and very delicate. The legs and antennae are 
colourless. The anal cleft is entirely absent, with the exception of a very short 
space posterior to the anal plates. Beyond this it is merely indicated as a line 
between the polygonal “cells” of the dorsum. Polygonal cells with straight sides, 
each with a very small transparent spot appearing as a point under the microscope. 
The space occupied by the anal plates is clear, faintly yellow, a striking contrast 
to the deep brown of the remainder of the dorsum when cleared. The inner face 
of the plates is straight ; the outer margin regularly curved to the tip giving them a 
broad heart-shape. (Fig. 243). The polygonal cells are fairly uniform in size to 
near the margin where they gradually disappear into the thin + structureless 
marginal area. Hye-spots distinct. 

The antennae are 8-jointed. Joints 2, 4, 6 and 7 sub-equal ; 1 a little shorter, 
3,5 and 8 longer. Measurements in w: (1) 30, (2) 37, (3) 47, (4) 30, (5) 50, (6) 30, 
(7) 27, (8) 47. 

Habitat: On upper side of leaf of avocado pear (Persea gratissima), from Mrs. 
Godwin, Durban, May 1916. 

Collection No.: 103. 

This species is remarkable for the hardness of the integument at maturity 
and te fact that the anal cleft is obsolete in the later stages. It differs from 
Hemilecanvum, however, in lacking the four perforate areas on the dorsum. 


12 CHAS. K. BRAIN. 


205. Saissetia kellyi, sp. n. 


Adult © almost circular, but rather wider than long (11:5 mm. by 13 mm.), flat, 
closely adherent to stem, deep chestnut in colour, with the dorsum matt, without 
ridges or lines of any kind, but flatly bounded to thin margin, often covered with 
dust, etc., so as to appear like a large flat blister on the bark. Younger specimens 
rather lighter in colour, with faint radiating ridges from the somewhat raised median 
area. 


The following particulars concerning antennae, legs, and microscopic characters 
in general, refer to insects measuring approximately 7 mm. long, i.e., before chitini- 
sation was fully complete. 

Antenna 9-jointed, range in w: (1) 48-54, (2) 85-102, (3) 129-153, (4) 75-85, 
(5) 58-65, (6) 105-116, (7) 44-51, (8) 34-48, (9) 74—79. 

Leg I: coxa 153, femur + trochanter 357, tibia 245, tarsus 153, claw 31. 


Anal plate about 320 ~ long. The margin has a single row of short, stout, conical 
spines. Stigmatic cleft with 3 gradually tapering spines, laterals about 50u, median 
about 136u. 

Around the body, just within the margin, is a series of complex, tubular glands, 
about 40 in number, distributed at almost regular intervals. Scattered over the 
integument, particularly near the margin, are numerous, short, stout, glandular 
hairs. The thin marginal area has the usual + straight, sided polygonal cells ; 
the median area has elongate oval cells much like those of Hemilecanium theobromae, 
Newstead. 

Habitat: On thick stems of Acacia melanoxylon, Pietermaritzburg, Natal ; 
collected by A. Kelly, June 1915. 

Collection No.: 118. 

The anal cleft is quite obsolete in mature specimens. This character, together 
with the two distinct types of cells on the dorsum and the 9-jointed antennae, 
reminds one of the type species of the genus Hemilecanium, Newst., but the absence 
of the four perforate areas preclude this species from that genus. 


206. Saissetia persimilis, Newst. (Plate 1., fig. 234). 

Lecanium (Saissetia) persvmile, Newst., Bull. Ent. Res. vu, p. 362, 1917. 

“Female, adult. Not differing appreciably in its external form, colour, and 
density of chitin from Lecaniwm (Saissetia) oleae (Bernard), but in two examples 
the dorsum was covered with a fine dusky-white, mealy secretion. The median 
longitudinal and two transverse ridges, forming roughly the letter H, well marked 
in two specimens, but absent in another. Anal cleft completely fused. Anal 
lobes attenuated, outer angle broadly rounded, inner edge much the longest ; apex 
bluntly pointed with one or two short spines. No stigmatic clefts; spines three, 
the central one slightly more than twice the length of the laterals. Antennae well 
developed, of eight segments. Legs rather slender; lower digitules very robust, 
incrassate proximally, dilated distally. Derm thickly studded with small, but 
well-defined, oval and translucent cells; these are much more crowded together 
at the margin and also larger.” (Newstead). 


THE COCCIDAE OF SOUTH AFRICA. 13 


Newstead states that Lecanium (Saissetia) sylvestru, Leon., has some similarity 
to persemile, but the former has 7-jointed antennae and three transverse ridges across 
the dorsum. 

Habitat : On Combretum, Muckleneuk, Pretoria ; collected in October 1914. 

Collection Nos.: 115 and 124. 


207. Saissetia subpatelliforme, Newst. (Plate 1., fig. 237). 

Lecamum (Sarssetia) subpatelliforme, Newst., Bull. Ent. Res. vu, p. 366, 1917. 

Adult © large, reaching 7 mm. long,5 mm. broad at the base and 3:5 mm. high, 
very convex but + conical, with the margin at the base slightly produced as a 
rounded ridge. The colour is dark castaneous to almost black, slightly shining, 
and without white flecks. There is no dorsal H, nor prominent ridges of any form 
(fig. 237). | 

Younger forms are lighter in colour, flat, disc-hke, with the margins slightly 
raised and rounded and the median area wrinkled. The anal cleft is apparently 
obsolete and the anal lobes inconspicuous. 

Antennae 8-jointed ; range in w: (1) 37-47, (2) 35-51, (3) 75-78, (4) 54-58, (5) 
41-54, (6) 24-27, (7) 17-20, (8) 51-58. 

Professor Newstead’s description, omitting figure references, is as follows : 

“Female, adult. Varying from broadly to narrowly ovate, centre generally 
highly gibbose; surface rather roughened or slightly rugose, often with widely 
separated patches of secretion, especially at the sides; dorsum (gibbosity) often 
shining. Derm cells of the median and sub-median areas small, ovate and widely 
separated, those near the margin much larger. Marginal spines of two kinds: 
(1) long and rather stout, dilated and divided on both sides ; (2) similar, but only about 
half the length of the long ones. Stigmatic clefts practically obsolete ; spines three, 
the laterals small, stout and pointed ; central one missing in all the preparations. 
Eyes well defined. Antennae of seven or eight segments (both forms present in 
one 2). Legs well developed ; tarsus almost equal in length to the tibia; lower 
digitules stout, upper pair normal. Anal lobes with the base and outer edge of 
equal length, approximately. Anal ringof eight hairs. Anal cleft varying in length 
from a little less than a third to one-third the length of the body. Length, 
3°8-5°2 mm.” 

Habitat: On stems of Cedrela toona, Salisbury, Southern Rhodesia; collected 
by Rupert Jack, June 1917. Also on stems of Syringa, Salisbury ; collected by 
Rupert Jack, February, 1909 (119a). 

Collection Nos.: 119 and 119a. 


208. Saissetia oculata, sp. n. 

Adult © elongate, 8 to 9 mm. long by 4 mm. wide, moderately flat, dull, buff to 
brown in colour, with the eye-spots and anal plates darker, almost black ; margin 
more chitin-brown, with a conspicuous fringe. Dorsum with rows (3) of hairs 
similar to fringe. Eyes distinct, almost circular, about 165 in diameter, with 
darker central prominence. Integument with faint oval cells, widely scattered. 
In the specimens examined these are most conspicuous around the anal lobes, 


14 CHAS. K. BRAIN. 


the point from which the chitinisation of the integument appears to spread in the 
LecaNtINAE. This suggests that, although the insects examined are adult, they 
may not be quite mature. 

Antennae of 8 segments, range in w: (1) 37-41, (2) 48-58, (3) 102-112, (4) 58-68. 
(5) 61-68, (6) 31-34, (7) 27-84, (8) 44-51. 

Leg I: coxa 88, femur + trochanter 238, tibia 170, tarsus 102, claw 24w. 

Margin with conspicuous fringe of long spines (136) with tips saw-like or finger- 
like. Dorsum with three longitudinal rows of long (143) spines. Anal plates long, 
tapering (280u). Stigmatic clefts inconspicuous with two very short spines and 
one longer (about half the length of marginal spines). 

Habitat: On grapevine, Durban; collected by C. P.v.d. Merwe, May 1917. 


Collection No. : 321. 


Genus Hemilecanium, Newstead. 


“ Adult female with the posterior extremity not cleft. Anal orifice placed near 
the middle of the dorsum and closed above with a pair of hinged plates as in 
Lecanium. Larva and nymph with a distinct anal cleft as in Lecanwum. Female 
in all stages with four large dorsal groups of compound spinnerets.” (Newstead). 


209. Hemilecanium theobromae, Newst. (Plate 1., fig. 238). 

Hemilecanium theobromae, Newst., Jl. Econ. Biol. ii, pt. 2, p. 39, 1908. 

Professor Newstead’s description, omitting figure references, is as follows :— 

‘Adult female broadly ovate or sub-circular; margins broadly flat, central 
area suddenly elevated, with strong irregular prominences. The whole of the 
dorsal area covered with a thin but dense layer of ochreous meal-like wax, but the 
prominences of the dorsum are generally denuded, apparently through abrasion, 
appearing through the secretion as shining, piceous, irregularities. Cuticle shining 
dark piceous on the central elevated area; dark castaneous and strongly rugose 
at the margins. Venter covered with a rather thick layer of white fibrous secretion. 
Anal cleft obsolete, but there is usually a faint emargination at the posterior 
extremity. Anal orifice placed just within the broad elevated dorsal zone at 
about one-third of the distance from the posterior margin. Derm cells 
of two distinct types; those of the dark central area narrowly elongat> 
with an apical pore, those of the pale broad margin of the ordinary polygona! 
type. There are also four groups of spinnerets; two towards the anterior 
and two towards the posterior extremity, each group composed of several hundre:i 
spinnerets forming well-defined dark chitinised areas. Antenna of nine segments, 
of which the third is the longest ; there are a few long hairs on the four terminal 
segments ; formula 3, 4 (5, 2, 9) 6, 1 (7,8). Legs highly chitinised ; slender but 
small compared with the size of the insect ; coxa almost equalling the length of 
the femur; digitules simple. Marginal spines with their broad bases suddenly 
contracted, each fitting into a well-defined socket, the latter being attached to a 
short subcutaneous tube. Stigmatic channels and spines absent. 

“ Length 13-15 mm. ; width 12-13 mm. 


er 


THE COCCIDAE OF SOUTH AFRICA. 15 


“Second stage female broadly ovate, slightly narrowed posteriorly, marginal 
spines continuous, resembling those in the adult. Antennae of seven segments, 
the third being the longest, the rest of the segments subequal in length. Legs 
scarcely longer than the antennae; coxa rather broad. Mentum uniarticulate. 
Groups of spinnerets occupying relatively the same position as in the adult, but 
there are only about 60-70 individual spinnerets in each group; they are also 
larger and more distinctly separated than in the adult. Derm in the region of the 
anal cleft finely squamose, with a large subcutaneous tube ; there is also a similar 
tube just within the margin opposite the anterior stigmata. 

“ Larva elongate; position of the compound spinnerets as in the adult and nymph. 
Mentum monomerous. Antennae of six joints, the third equalling the length of 
the fourth, fifth and sixth together. Marginal spines forming a continuous series.” 

Habitat : On tree euphorbia, East London, and stems of oleander, Capetown 
(Fuller, 1898). 

Collection No. : 107. 


Genus Aclerda, Sign. 
Adult 2 naked, a mere sac containing eggs at maturity, with normal mouth 
parts, but legs and antennae absent. 
Larva greatly elongated, with parallel sides. 


210. Aclerda digitata (CklI.), 

Pseudolecanium digitatum, Ckll., Ann. Mag. Nat. Hist. (7) ix, p. 24, 1902. 

Aclerda digitata (Ck) Fernald, Catalogue, p. 210, 1903. 

Professor Cockerell’s description is as follows :— 

“ , pyriform, with the hind end pointed ; about 2} mm. long; dark ferruginous, 
shiny, producing some cottony material. End of abdomen strongly chitinized, 
with the form usual in the genus. The diagnostic characters, as in all species of 
the genus, are derived mainly from the larva, the female being a mere bag of eggs. 

“ Larva extremely long and narrow, length 600, breadth 160. Anterior extremity 
truncate, crenulate, with six short blunt, finger-tip-like spines; a row of fifteen 
blunt spines down each side of body; no dorsal spines; legs well developed, 
anterior tibia 60, its tarsus 30u long; the tibia has a constriction about 27 from 
base, making it look almost 2-jointed, this being more or less apparent on all the 
legs ; tarsal digitules long, with small knobs; claw-digitules shorter, filiform, 
knobbed. Antennae 24 apart at base and 30u from anterior end of head; 6- 
jointed, joints measuring: (1) 18, (2) 12, (3) 27-33, (4) 20, (5) 21, (6) 30. Anal 
ring small, circular, without bristles on its margin, but posterior to it are set four 
bristles, of which the inner two are shortest ; anterior to anal ring is a row of four 
finger-like blunt spines ; on each side is a caudal bristle, about 190 long, and beyond 
each of these a pair of finger-like spines; anterior to and a little laterad of each 
caudal bristle is a small round gland. The last two segments have each a pair 
of bristles on the ventral surface, those on the last being twice as long as those on 
the penultimate segment.” 

Habitat : On grass, Natal; collected by C. Fuller, 1901. This species has not 
been found again. 

Collection No. : 134. 


16 CHAS. K. BRAIN. 


Genus Allopulvinaria, nov. 

Adult 2 naked above, very convex and distinctly segmented. Lower surface 
concave, filled with a compact powdery mass of wax which projects around the 
margins. Antennae somewhat rudimentary, of 4, 5 or 6 segments. Legs well 
developed. Anal ring with numerous (12?) hairs. 

Larva elongate and narrow ; anal tubercles produced, each with one long seta. 


Type, A. subterranea, sp. n. 


211. Allopulvinaria subterranea, sp. n. (Plate vi, fig. 246). 

Larva active, pale pink, eyes pigmented, body elongate and narrow. Anal 
tubercles long, each with one long seta. 

One-fourth grown 2 with 4-jointed antennae, and marginal hairs as in Lecaniwm. 

Adult @ viviparous, 7 mm. long, 45 mm. broad and 4 mm. high. Dorsum 
roundly arched ; segmentation distinct, the segments roundly ridged. 

The body is red, faintly flecked with white wax. Underside flatly concave, 
abdominal portion plainly segmented. Legs small, brown. Anal plates small, 
brown, from between which protrudes a double fine pencil of white wax. Lower 
concavity of body filled with cushion of white wax which extends slightly beyond 
the margin of the body; this is not cottony but compact. No eggs were seen, 
but larvae were present in the white waxy cushion and in the body of the Q. 

Antennae 4, 5, or 6-jointed, usually 5. Anal ring with numerous (12?) hairs. 

Anal plates raised, appearing excavate so as more or less to enclose the anal 
tube from the sides as well as above. Anal groove closed. Integument with 
numerous simple glands and short acute spines scattered over the surface; the 
former especially noticeable in transverse series across the segments. 

The following measurements may be useful for comparison. 


Antennae: (qa) 4-jointed type :— 
I 


W I IV. 
(a) 40 27 61 34u 
(b) 34 40 65 b4u 


(b) 5-jointed series, range in yu :—(1) 34-45, (2) 3440, (3) 51-68, (4) 15-34, (5) 
23-40; (c) 6-jointed form, e.g. (1) 41, (2) 34, (3) 61, (4) 17, (5) 10, (6) 20. 

Leg II: coxa 81 by 108, femur + trochanter 150, tibia 74, tarsus 74. 

Habitat: On stems of “ quick” grass, near bank of stream ; collected by T. L. 
Watermeyer, Jonkershoek, Stellenbosch, C.P., July 1917. 

Collection No.: 138. 


Genus Protopulvinaria, Ckll. 

The few species which have been described in this genus are similar to Lecanwum 
in their early stages, but may be readily distinguished at maturity by the small 
white ovisac which is secreted beneath the adult 9. In the normal condition this 
is but slightly larger than the body of the female and protrudes as a white ring around 
the margins of the body (fig. 248). 

This genus is included in Pulvinaria by Mrs. Fernald in her Catalogue (p. 128), 
but it would appear that its correct position is between Lecanium and Pulvinaria. 


THE COCCIDAE OF SOUTH AFRICA. 17 


212. Protopulvinaria piriformis (Ckll.), Lefroy. (Plate il, fig. 244; iv, fig. 248.) 
Pulvinaria (Protopulvinaria) pyriformis, CkIl., Jl. Trin. Nat. Club, i, p. 309, 1894. 
Pulvinaria (Protopulvinaria) pyrvformis, Ckll., Jl. Trin. Nat. Club, ii, p. 307, 1896. 
Pulvinaria newsteadi, Leon., Riv. Pat. Veg. vi, p. 279, 1898. 

Pulvinaria pyriformis, Ckll., Psyche, vii, p. 311, 1899. 

Protopulvinaria pyriformas, Lefroy, Scale Ins. Lesser Antilles, p. 43, 1901. 

Pulvinaria pyriformis, Fernald, Cat., p. 138, 1902. 

Pulvinaria plana, Ldgr., Jahrb. Hamb. Wiss. Anst. xxxii, p. 34, 1911. 

Protopulvinaria piriformis, Ldgr., Die Schildliuse, p. 199, 1912. 

Ovisac slightly larger than the body of the adult 9, from which it extends as a 
uniform ring around the margins (fig. 348). Ova pale greenish white, regularly oval. 

Adult 2 about 3 mm. long and about as broad slightly behind the middle ; 
broadly rounded behind, suddenly narrowing in front so as to be short pear-shaped 
or drop-shaped. The margins of the body are flat ; the central part is flatly rounded. 

In living specimens the general body colour is yellowish or pinkish with the margins 

reddish brown. The subdorsal area is irregularly suffused with mauve or violet. 

The anal cleft is deep, extending almost to the centre of the body, but the sides are 

in close proximity, so that the posterior margin in most cases appears uniformly 

broadly rounded. The anal plates are very long and very narrow and are brownish 
in colour. Dead, dry specimens are pale to deep brown according to age. When 
stained, cleared and mounted the following characters may be noted :— 

Antennae 7-jointed ; range in w: (1) 27-31, (2) 37-44, (3) 44-51, (4) 51-54, (5) 

20-24, (6) 20-24, (7) 51-54. 

Leg I: coxa 75, femur + trochanter 170, tibia 109, tarsus 54, claw 20wu. 
Leg II: coxa 99, femur + trochanter 180, tibia 112, tarsus 68, claw 24. 


Anal plates very long (470) and narrow (fig. 244). 

Marginal spines short, thin, with ends deeply branched, or occasionally bifid 
from base, in a close-set series. Stigmatic spines 3, laterals short and acute, median 
two or three times as long. Young form, about 1:2 mm. long, oval, translucent, 
appearing faintly greenish yellow, with six deep purple lines radiating from median 
area to margin, the four posterior being nearer together than the anterior two. 

Habitat : On undersides of leaves of avocado pear (Persea gratissima) ; collected 
by A. Kelly, Pietermaritzburg, April 1916. 

Collection No.: 77. 


Genus Pulvinaria, Tare. 

‘* Adult female resembling Lecanium in the early stages and until the commence- 
ment of oviposition, when a loose cottony ovisac is secreted from below the posterior 
extremity of the insect, for the protection of the eggs. The body of the insect is 
tilted up during the formation of the ovisac, and often becomes much shrivelled 
and distorted, finally remaining as a small shapeless scale at the anterior extremity 
of the mass of ovisac. It is sometimes rendered still more inconspicuous by the 
partial overlapping of the secretionary matter, but is never completely enveloped. 
In all purely structural characters there is nothing to distinguish species of this genus 

(681) | B 


18 CHAS. K. BRAIN. 


from those of Lecaniwm, so much so that, until the period of oviposition, it would be 
impossible to determine whether an individual should be placed in the one or the 
other genus.” (Green). 


213. Pulvinaria aristolochiae, Newstead. (Plate ui, fig. 242). 
Pulvinaria aristolochiae, Newst., Bull. Ent. Res. vin, p. 19, 1917. 


Adult 9, immediately prior to forming the ovisac, about 8 mm. long and 4:5 mm. 
broad, with thin margins and the dorsum rising to a distinct central prominence : 
colour brown, with darker markings. The margin is supplied with a short fringe of 
thin pale hairs ; from this to the raised centre extend a number of dark lines. 

Male puparium about 3 mm. long, moderately convex, dull white, not glassy, 
with a distinct transverse ridge about the middle. Two white filaments protrude 
from the posterior extremity some days prior to the emergence of the adult ¢. 
Adults were emerging in July 1916. 

Antennae 8-jointed, range in « :—(1) 57-75, (2) 68-75, (3) 187-190, (4) 95-102, 
(5) 78-85, (6) 34-47, (7) 24-34, (8) 41-44. 

Leg I: coxa 185, femur + trochanter 400, tibia 262, tarsus 123, claw 34u. 

Anal plate about 250u long. Marginal spines short (27-35), many appearing 
truncate. 

As Professor Newstead did not have the opportunity of examining living material 
the above particulars are given to supplement his description which is as follows :— 

“Female, adult. More or less cordate in outline and rather flattened ; dorsum 
almost completely covered with a well-defined layer of flake-like wax which varies 
in colour from dirty grey to greyish brown. Antennae of eight segments ; 3rd, 
4th and 5th unusually long, the two first-named swollen distally ; 3rd about twice 
the length of the 4th ; three long hairs on 2nd, one of which lies (in three examples) 
close up to the succeeding segment ; there is also a long distal hair on the 3rd ; 
two on the 5th; a single spine on the 6th, 7th and 8th, the last-named also with a 
few very short hairs. Legs stout, long; tarsus relatively very short, less than 
one-third the length of the tibia ; lower digitules long and very broadly spathuliform. 
Marginal spines stout, pointed, and placed rather close together ; stigmatic spines 
broken away in all the specimens, their points of attachment being continuous with 
the marginal series. Submarginal pores very large, continuous, but rather widely 
separated. Anal lobes with four stout spines near the apex, on the inner edge. 
Anal cleft short, usually a little less than one-sixth the entire length of the body. 
Anal ring with eight hairs. No derm cells present; but there are numerous 
circular spinnerets (? ventral), each having an inner concentric ring. Length 
57-76 mm. 

“ Ovisac pure white and closely felted, long and generally tortuous. Length 
10-20 mm.” 

This insect shows great similarity to P. jacksoni, Newstead. Both have remark- 
ably long joint 3 to the antennae, which in jacksoni is almost 120y, in aristolochiae 
about 180u. Further both produce extremely long ovisacs. 

Habitat: On stems of Erythrina caffra Thunb., (Kaffir-boom), Natal Coast. 

Collection Nos.: 79 and 84. 


THE COCCIDAE OF SOUTH AFRICA. 19 


214. Pulvinaria floccifera (Westw.). 

Coccus floccifera, Westw., Gard. Chron., p. 308, 1870. 

Pulvinarva brassicae, Ckll., Can. Ent. xxvu, p. 135, 1895. 

Pulvinaria floccifera, Green, Ent. Mon. Mag. xxxiu, p. 72, 1897. 

Pulvinaria pharae, Lull, Ent. News, x, p. 237, 1899. 

Pulvinaria floccosa, Newst., Jl. Roy. Hor. Soc. xxii, p. 26, 1900. 

Completed ovisac about 6 mm. long and 3 broad, with parallel sides and rounded 
extremity ; pink in colour, from the numerous ova which are of that colour; the 
small amount of white cottony matter which is incorporated in the mass is incon- 
spicuous owing to the large number of eggs. 

©, young stage: about 2 mm. long, translucent greenish yellow, with the median 
dorsal area suffused with a pinkish or brownish tint. 

Adult 2 about 3mm. long and almost as broad, moderately convex. The 
margins are pale translucent greenish yellow, the median dorsal area is reddish 
brown and the whole dorsum is more or less wrinkled. The crests of the ridges, 
especially those running to the margin, blackish brown. Anal plates small, of 
the same colour as the dorsal area or slightly paler. When the ovisac is almost 
complete, the body of the Q has shrunken to such an extent that the brownish 
part of the dorsum is not conspicuous, while the black crests of the ridges are 
close together and thus give the insect a much darker appearance. 

Antennae 8-jointed, range in w: (1) 34-88, (2) 44-51, (3) 65-72, (4) 34-44, (5) 
34-41, (6) 24-27, (7) 24, (8) 44. 

Leg I: coxa 78, femur -++ trochanter 190, tibia 163, tarsus 102, claw 24w. 

Upper digitules very long, slender, with spherical knobs. Lower digitules stout, 
normal. Anal plate about 145u long. Just in front of the anal plates there are a 
few scattered, “rosette” glands and two pairs of long (108) hairs. Marginal 
spines about 50u long and 100u apart, slender, often with finely divided tips, alter- 
nating with a submarginal row of similar hairs of about half the size. Stigmatic 
spines comparatively short and stout, with the outer ends tapering to blunt points. 
From these to the spiracles extends a double row of small rosette glands. 

Puparium of 3 of the usual type with the segments of the plates very distinct. 
The whole puparium is opaque white, appearing finely striate in a longitudinal 
direction. 

Habitat: On a native plant (Solanwm sp. ) Durban ; collected by C. P. v. d. 
Merwe, July 1916. 

Collection No. : 82. 

A very large percentage of the young forms (about 1:5 mm. long) are very 
convex and shiny black, obvious!y parasitized. 


215. Pulvinaria jacksoni, Newst. 

Pulvinaria jacksoni, Newstead, Jl. Econ. Biol. i, p. 155, 1908. 

Professor Newstead’s description is as follows :— 

“ Ovisac from two and a half to seven times the length of the female ; breadth 
equal to the width of the insect ; low convex, sides parallel ; closely felted, tough 
and web-like in texture. Length 16°75-42 mm. 

‘681) Rn? 


20 CHAS. K. BRAIN. 


‘* Adult female—Dried examples sienna brown or dull ochreous; the pale examples 
with yellowish mottlings indicating the position of the derm-cells. Form normal. 
Antennae of eight segments, the third being much the longest; last four subequal ; 
there is a very long hair on the third and fifth, the one on the former the longest. 
Legs stout ; trochanter about one-third the length of the femur, with a very long 
apical hair; digitules to claws and tarsi normal. Derm with very small incon- 
spicuous glands. Marginal spines short, closely set, truncate, the ends often 
notched. Stigmatic clefts with pointed spines and two large tubercles. 

“Male puparium, much stained with brown, normal in shape, but thick and un- 
usually wax-like in appearance; dorsal plate with one large central, white, wax- 
like felted spot and several brownish tubercles immediately beyond it. Length 2°25 
mm.” 

The following measurements are added for comparison with the other South 
African species :— 

Antennae 8-jointed ; range in w: (1) 48-54, (2) 54-68, (3) 102-116, (4) 68-75, 
(5) 61, (6) 31-34, (7) 31-34, (8) 37-41. 

Leg I: coxa 139, femur + trochanter 354, tibia 230, tarsus 108, claw 27. 

Anal plate about 240u long. 

Habitat: On Trichitla sp., Durban ; collected by A. Kelly, May 1918. 

Collection No.: 79a. 


216. Pulvinaria lepida, sp. n. 

Adult °° with ovisacs clustered on stems and leaves of grass. 

Adult 9 averaging 2 mm. long by 1°3 mm. broad, moderately arched, ight brown 
in colour, dorsum with transverse wrinkles; marginal area flecked with white 
wax. Ovisac white, elevated, extending behind the insect for about the length 
of 2 body, faintly fluted on upper surface. 

Numerous 9° parasitised. Mounted specimens contain numbers of well 
developed embryos. 

Antennae variable, of 7 or 8 segments; terminal joimt appearing truncate ; 
range in wu :—T-jointed : (1) 34-37, (2) 37-48, (3) 54-65, (4) 70-82, (5) 24-27, (6) 
24-27, (7) 37-41; 8-jointed : (1) 34-87, (2) 44-48, (3) 51-58, (4) 34-41, (5) 44-48, 
(6) 24-27, (7) 24-27, (8) 37-41. 

Leg I: coxa 78, femur + trochanter 187, tibia 153, tarsus 102, claw 24y. 

Anal plate about 153 long, with 5 comparatively short hairs at the apex. Mar- 
ginal spines in a single row, simple, pointed, about 41u long and 20-30 apart, 
straight or only slightly curved. Stigmatic spines: laterals short, acute ; median 
stouter, longer, blunt. 

Halitat: On stems and leaves of common veld grass, Standerton and Pretoria 
(K. Munro). 

Collection Nos. : 139 and 328. 


217. Pulvinaria mesembryanthemi (Vallot), Sign. (Plate i, fig. 235). 
Pulvinaria mesembryanthemi, Vall., Bull. de Ferussac, xxu, p. 469, 1830. 
Calypticus mesembryanthemi, Costa, Ann. Acad. Asp. Naples, p. 273, 1844. 
Pulvinaria biplicata, Targ., Catalogue, p. 34, 1869. 

Pulvinaria mesembryanthema, Sign., Ann. Soc. Ent. Fr. (5) ii, p. 39, 1873. 


THE COCCIDAE OF SOUTH AFRICA. 21 


Adult 9.to time of forming ovisac, and younger stages, green, of the same tint 
as the fleshy leaf of Mesembryanthemum edule, its most common food-plant ; 
attaining a length of 5 mm. and an almost equal breadth, moderately convex, 
dorsum smooth. 

As the ovisac is produced the body of the 2 becomes yellowish and later yellowish 
brown, much shrunken, with four transverse ridges and, ultimately, contorted 
or bent backward (fig. 235). 

Antennae 8-jointed; range in w: (1) 48-55, (2) 48-54, (3) 75-85, (4) 44-58, 
(5) 27-37, (6) 24-34, (7) 27-31, (8) 46-51. 

Leg I: coxa 122, femur + trochanter 290, tibia 211, tarsus 126, claw 34u. 

Anal lobes approximately 165 long. 

This insect is common throughout South Africa on Mesembryanthemum spp., 
especially M. edule, Linn. It becomes so numerous in some seasons that it kills 
patches of this plant when grown in parks, etc. This was the case at the Eastern 
Sports Grounds, Pretoria, in November 1914. 

The 3g emerged from the next generation about Christmas 1914 and the following 
particulars are given from fresh material obtained at that time :— 

Male test, transparent, white, glassy, about 1‘5 mm. long, margins depressed, 
central plate raised, slightly keeled. 

Body, legs and antennae dark brown. Wings broadly rounded, iridescent, dorsal 
sclerites and eyes shiny black. Two caudal waxy filaments, white, as in Pseudo- 
coccus, length equal to head + body without antennae. 

Length of head + body 1:17 mm. (without genital spike) ; length of wing 1:0 mm.; 
width of wing 0°5 mm.; length of antenna 0°84 mm.; length of genital spike 
0°27 mm. ; length of caudal setae 1:0 mm. 

Antennae 10-jointed, the segments measuring: (1) 34, (2) 48, (3) 51, (4) 180, 
(5) 112, (6) 105, (7) 85, (8) 68, (9) 61, (10) 74. 

Habitat: On Mesembryanthemum spp., chiefly M. edule, Linn., throughout the 
Union. 

Collection No: 78. 


218. Pulvinaria psidii, Mask. 

Pulvinaria psidu, Mask., N.Z. Trans. xxv, p. 223, 1892. 

Adult 9 pale transparent yellow with irregular black markings; 2 mm. long 
and 1:7 mm. broad. The ovisac is white, as wide as the body, always flat, attaining 
twice the length of the body. Ova pale yellow. 

Larvae pale yellow, hatching when received, 16th November 1914. 

Adult 9 flatly convex with slightly raised median keel. Anal plates pale brown. 

Antennae 8-jointed; range in ww: (1) 24-34, (2) 37-44, (3) 51-58, (4) 27-34, 
(5) 27-34, (6) 20-24, (7) 14-20, (8) 41-44. 

Leg I: coxa 68, femur + trochanter 190, tibia 145, tarsus 85, claw 271. 

Anal plate about 130u long. Margin with a row of thin spines about 34, long, 
tips divided but not dilated. These spines are set much closer together on the 
anterior part of the body. Integument clear, with a few simple glands with short 


29, CHAS. K. BRAIN. 


spines and a few long hairs near the insertion of the antennae. Stigmatic spines ; 
laterals short, conical ; median longer (51y). 

Habitat : On guava, Botanic Gardens, Durban ; collected by A. Kelly, November 
1914. 

Collection No.: 81 


Genus Ceronema, Mask. 

Maskell’s original diagnosis of this genus, as given in the Transactions of the New 
Zealand Institute, p. 55, 1894, is as follows :— 

“ Female insects in the adult stage covered wholly or partially by tests of threads 
more or less closely woven, neither glassy nor felted, never forming homogeneous 
plates. No fringe. Form of insect Lecanid, with normal cleft and lobes. 

“ Larva Lecanid, showing cleft and lobes. 

““ Male pupa covered by a glassy test of normal Lecanid form, composed of plates 
more or less homogeneous.” 


219. Geronema mobilis, sp. n. 

Adult 9° about 3 mm. long, broad oval in outline, flat, with the dorsum almost 
entirely covered with long, white, waxy filaments. Around the margin the filaments 
are long and coarse, but those on the dorsum are short and fine and + curly and 
densely matted and appear almost felted. The colour of the insect, beneath the 
secretion, is caramel brown. 

When cleared and mounted the body is broad oval ; integument thin and trans- 
parent with numerous scattered, small, simple gland openings. Margin very 
closely set with a very short fringe of marginal spines, which are only about 20u 
long, thin, linear, two or three, usually two, arising from the same pore; they are 
unusually close together, averaging about 14u apart. Stigmatic clefts shallow, 
but distinct because of their chitinised sides; each cleft with two or three curved, 
blunt processes which are about as long as the marginal spines. 

Antennae 7-jointed; range in w: (1) 27-37, (2) 41-51, (3) 34-54, (4) 68-78, 
(5) 27-31, (6) 17-27, (7) 41-44. Joints 2 and 4 each with two long setae, 5 
with one, 6 with two and 7 with eight. 

Leg I: coxa 75, femur + trochanter 187, tibia 105, tarsus 68, claw 24u. 

Anal plate about 140u long with | apical and 3 subapical spines. 

Habitat: On leaves of common native bush (Celastrus cordata, E.M.) (Celastri- 
neae) ; collected by C. Fuller, Illovo River, Natal, August 1916. 

Collection No.: 97. 

The adult 2 moves about quite readily. A number of mature specimens 
were put on one side when received with a view to photographing them, but 
next day, when the leaves had slightly dried, the insects had all crawled off and 
so disarranged the waxy filaments. 


Genus Lichtensia, Sign. 
Adult 9 oval or suboval, like a Pulvinaria, except that the secretionary sac covers 
the whole body with the exception of the cephalic end, which is usually more or less 
exposed. Antennae 8-jointed. 


THE COCCIDAE OF SOUTH AFRICA. 23 


220. Lichtensia asparagi, sp. n. 

Female ovisac about 4:5 mm. long, elongate, narrow and very convex, sordid 
white in colour, thin but dense, with the anterior extremity open, exposing the 
front portion of the dry female. 

Male puparium moderately large, elongate, very convex, with perpendicular 
sides, thin, semi-opaque, dorsum flatly rounded, entirely without median plate. 
There is a faint indication of sublateral lines, otherwise the surfaceis uniformly stippled. 

Adult 9, mounted, broad oval, about 4 mm. long, with the two extremities 
about equally rounded ; anterior extremity not produced. 

The integument appears remarkably free from conspicuous glands or hairs. 
Margin with a single scattered row of short sharp curved spines. Stigmatic spines 
stout, somewhat curved, with rounded ends. 

Antennae 8-jointed, in one antenna examined 9-jointed, with the following range 
in w: (1) 35-48, (2) 57-60, (3) 68-78, (4) 51-71, (5) 40-61, (6) 30-37, (7) 25-34, 
(8) 30-38. 

Leg I: coxa 88, femur + trochanter 228, tibia 190, tarsus 102, claw 24. 

Anal plate about 185 long, with longitudinal folds. 

Habitat: On Asparagus capensis, L. (Liliaceke), Eastern Cape Province. 

Collection No. : 131. 


Genus Filippia, Targ. 
Female flat, oval, legs and antennae normal. Side of body and dorsum with 
numerous tubular glands. Female, at maturity, entirely enclosed in a flat felted 
sae which, after oviposition, serves as an ovisac. 


221. Filippia chilianthi, sp. n. 

Adult 9 entirely covered with a dense, closely felted layer of white secretion, 
about 6 mm. long and 3 mm. broad, + regularly oval, with the dorsum moderately 
convex. ‘This secretionary layer is secreted slowly from all the upper parts of the 
body, the glands concerned being scattered. This is not the case in the Pszupo- 
COCCIN AE, in which the ovisac is formed by secretionary glands in the posterior and 
marginal portion of the body only. Little or no secretion appears on the ventral 
surface, so that the leaf or twig forms the floor of the sac; the marginal secretion 
becomes attached and entirely encloses the 2. As oviposition proceeds the body 
shrinks away from the posterior end of the sac which becomes almost filled with ova. 
At death the dry shrivelled body of the 9 occupies a small portion at the anterior end. 

Prior to secreting the ovisac the 2 is from 4 to 5mm. long, and rather more 
than half as wide. The sides are almost parallel for more than half the length of 
the body, the posterior end appearing broadly rounded, and the anterior end more 
pointed. In colour it appears greyish brown with a lighter median, flatly rounded, 
keel. When seen under a hand-lens, however, it is noticed that the ground-colour 
of the body is yellowish, and semi-transparent, speckled with black. The anal cleft 
is short, with the sides closely adjacent. The anal plates are small and brown. The 
lower surface is slightly concave, greasy-yellowish in colour, the black speckling of 
the dorsum only showing faintly through at the extreme margins. The stigmatic 
bands are represented by two short faint white lines on each side. 


24 CHAS. K. BRAIN. 


At a younger stage—when about 2°5 mm. long—the body is red-brown, rugose at 
the edges, with the median keel not prominent but smoother than the remainder 
of the body. 

The antennae are 8-jointed; range in uw: (1) 31-45, (2) 50-62 (3) 95-108, 
(4) 54-65, (5) 32-44, (6) 27-34, (7) 27-34, (8) 31-37. 

Leg I: coxa 102, femur + trochanter 245, tibia 165, tarsus 85, claw 20u. 

Tarsal digitules moderately long (70x), slender hairs with their distal ends slightly 
clubbed. Lower digitules short, broad. Marginal spines short (27), truncate. 
Anal plate about 155u long. Stigmatic cleft with three comparatively slender 
spines, laterals about 34, median similar but longer (51,1). 

Male puparium about 2 mm. long and 1 mm. wide, dark red when containing the 
insect ; white and glassy when empty. The divisions of the test are very prominent, 
opaque white ; the remainder being + wrinkled and almost hyaline. 

Adult 3g were emerging in large numbers when the material was received, 
5th April 1916. 

Copulation was observed in a number of cases, the 9 in each case measuring about 
4mm. in length. The body of the male was curved downward admitting the sheath 
into the cleft of the Q. During this process the two long waxy filaments of the J 
stood erect. 

The adult ¢ is about 1:0 mm. long, rich red-brown in colour, with two long waxy 
caudal filaments about half as long again as the head and body combined, without 
the antennae. The body, legs, and antennae are all red-brown. The head is slightly 
darker. The scutellum is polished at the edges, with a sunken, matt, central patch. 
The wings are hyaline, matt, extremely iridescent, with a strong sub-costal, cochineal 
red band. 


When mounted, the following measurements may be taken as an average :— 


Head and body, without antennae .. se mn os ul taba pe 
Antennae... nf ai ae fe vf .. 0847 mm. 
Genital spike .. sys i oh Ss - Je fe. em. 
Wings : length 1-2 mm., width Hs she be 3} OG, ain 
Caudal filaments 20 2 ft £20087, om: 


Habitat: On a native shrub ; collected by C. Fuller, [lovo River, Natal, April 
1916. On leaves of Chilianthus oleaceus, Burch (Loganiaceae), Bloemfontein, 
0. F.S.; collected by J. C. Faure, April 1916. 

Collection Nos. : 127 and 128. 

This insect, with the ovisac completed, is similar in appearance to F. oleae 
(Costa) but, according to Signoret’s description of that species, the adult has 
6-jointed antennae. 


222. Filippia carissae, sp.n. (Plate i, fig. 236). 

This species differs from F’. africana in the following respects :— 

The ovisac is a little larger and generally less compact ; often split around the 
edges (fig. 236). The antennae are more variable, 7 or 8-jointed, range in wu :— 


THE COCCIDAE OF SOUTH AFRICA. Zo 


(a) 8-jointed: (1) 41-54, (2) 51-54, (3) 102-119, (4) 68, (5) 37-44, (6) 34, (7) 24-27, 
(8) 34-37; (b) a specimen with 7-jointed antennae gave the following measure- 
ments :— 

1. 54 68 105 122 34 27 41 
il. rt aR 109 68 51 37 68 


Leg I: coxa 95, femur + trochanter 240, tibia 204, tarsus 88, claw 20u. 

Anal plate 176u, rather elongate, with one or two long spines at apex. Margin 
with a single row of + plate-like spines, narrow at the base and gradually 
broadening at the distal ends. Stigmatic clefts somewhat sunken, with three spurs 
of moderate length. 

Habitat: On leaves of Carissa grandiflora, A. DC. (Apocynaceae), Natal Coast ; 
common. 

Collection No. : 129. 


Genus Conofilippia, nov. 

Female Lecaniid, flat, with well developed legs and antennae; dorsum with 
numerous sharp spines; margin with slender spines and stigmatic cleft similar 
to Lecanium. At maturity the insect is entirely enclosed in a dense felted sac, which 
is elevated in the form of a high cowl. Antennae 7-jointed. 


Type, C. subterranea, sp. n. 


223. Conofilippia subterranea, sp.n. (Plate iui, fig. 245). 

Adult °° enclosed in conical felted tests on the roots of a native shrub. Test 
about 6 mm. long, 5 mm. broad and 5 mm. high (figs. 245a, 6). 

Adult © flat, about 5°5 mm. long, 4mm. wide, smooth, glossy, pink to red in 
colour, brown and wrinkled when dry, on a thick mat of dense powdery material 
(the floor of the test) which is buff to yellow-brown in colour and dusted with white 
wax beneath the insect. 

In one or two cases the top of the test shows a circular orifice, but this does not 
appear to be normal and looks as though it had been made for the exit of some 
Hymenopterous parasite. 

Largest 2, when mounted, about 6 mm. long, and 4:2 mm. wide. Integument 
clear, with a number of simple glands which are particularly abundant near the 
margin, and with numerous short (17) sharp spines. The margin is thickly set 
with a compound series of slender spines about 24 long, which are somewhat curved 
near their tips. Stigmatic clefts with three blunt spurs, laterals about 27, median 
about 371. : 

Antennae 7-jointed; range in mw: (1) 35-40, (2) 25-34, (3) 40-47, (4) 37-42, 
(5) 34-44, (6) 17-30, (7) 37-51. In one case joints 6 and 7 were united, forming a 
segment 64 long. 

Leg I: coxa 102, femur + trochanter 220, tibia 170, tarsus 100, claw 40u. The 
junction of the tibia and tarsus is swollen and suddenly narrows towards the distal 
half of the tarsus. 

The anal lobes are rounded, about 230m long. 


26 CHAS. K. BRAIN. 


Habitat: On roots of native shrub with red stems; collected at De Wildt, 
Pretoria District, Transvaal, by Claude Fuller, May 1915. The drawings (figs. 
245-b) were kindly made from living material by the collector. 

Collection No.: 137. 


Genus Geroplastes, Gray. 

Adult 2 completely enveloped in a more or less dense covering of wax ; no marginal 
fringe or radiating processes. Sometimes the waxy covering exhibits a variety 
of arrangement in the form of definite plates or plaques. In other species the 
covering is uniform in texture and is then most often soft, and contains a large per- 
centage of watery fluid. On removing the wax a caudal prominence is generally 
visible. Legs and antennae present, well developed. 


224. Geroplastes bipartitus, Newst. 

Ceroplastes bipartitus, Newst., Bull. Ent. Res. viu, 1, p. 25, 1917. 

“Female test. Colour of dried specimens very like pale dirty beeswax. In 
the young adults the test is broadly oval, somewhat hemispherical and divided 
into nine plates: three bilateral, one cephalic, one anal and one dorsal, the last- 
named with a conspicuous dark brown or blackish oval spot, with a central elongated 
patch of pure white wax; the nuclear spots to the lateral plates are smaller and 
generally much less conspicuous than the dorsal one. Margin over the stigmatic 
areas with a pair of laterally compressed and somewhat disc-shaped extensions,, 
each extension carrying on its edge a narrow strip of opaque white wax, the tip 
of which sometimes reaches the dark nuclear spot of the lateral thoracic plate. 
In very old examples the test has increased in thickness considerably, but this has 
been so much damaged in transit as to render it useless for descriptive purposes ; 
however, one can trace the curious marginal extensions, which are somewhat like a 
narrow-waisted and distorted bobbin, or the toy used in the once popular game 
‘diabolo.’ Average length of young adults, 3 mm.; height 1-6-2 mm.; average 
length of old adults, 6 mm. ; height doubtful. 

“Female adult. Denuded of wax, hemispherical; caudal process very long, 
varying in length from one-half to a little less than one-half the length of the 
remaining portion of the insect. Submarginal tubercles small, but generally 
clearly defined ; one cephalic and three bilateral, the two over the stigmata slightly 
more pronounced than the rest. When examined under a high power lens, by 
transmitted light, these tubercles are seen to be traversed by clear cell-like tracts. 
forming an irregular reticulated pattern. It is reasonable to assume, therefore, 
that these may be the special set of glands which secrete the nuclear spots in the 
centre of the plates in the test. Derm relatively thin, but strongly chitinised. 
Pores minute, separated over a large portion of the dorsum by slightly varying 
distances equalling the length of one of the short segments of the antennae or two of 
them together. Stigmatic clefts relatively shallow, but very clearly defined ; spines. 
short, obconical, those at the extreme margin very minute and stud-like. Marginal 
spines or hairs not traceable. Antennae of six segments, the 3rd longer than the 
last three together. Legs normal. Length of denuded female, inclusive of the: 
caudal process, 4°5-4°6 mm. ; length of caudal process, 13-15 mm. 


TNE COCCIDAE OF SOUTH AFRICA. 27 


“Male puparium. Consisting of two distinct parts ; the lower half boat-shaped, 
and of a glassy vesicular texture, as in those typical of the genus Lecaniwmn ; the 
upper portion opaque, low, convex, and of a dirty beeswax colour, with nine narrowly 
rectangular, submarginal patches of snow-white secretion. Anal cleft apparently 
obsolete. On the emergence of the male the whole of the upper portion falls away, 
leaving the ventral half attached to the food-plant. The line of cleavage between 
the upper and lower portion is clearly defined in those puparia from which the 
imprisoned male has not escaped. Length 1:6 mm. 

‘South Africa, 1914 (de Charmoy).” (Newstead.) 

This species is apparently not represented in the collection of this Division. 


225. Ceroplastes candela, Ckll. & King. 

Ceroplastes candela, Ckll. & King, The Entom. xxxv, p. 113, 1902. 

“©. Long 23, lat. 34, alt. 44; dark red-brown, elevated, with vertical s:des. 
Caudal horn a prominent stout spine, hardly $ mm. long, placed nearer the top 
of the scale than the base. Dorsum smooth and shining, with only a very small 
central raised line. Sides of insects with vertical stripes of dense secretion ; no 
wax, except that composing these stripes, between the insects, which are densely 
crowded together, their vertical sides contiguous. They rest on a thin substratum 
of wax, and are covered above with yellowish-white wax, about 1 mm. thick. The 
outlines of the insects are vaguely marked on the surface of the covering wax by a 
brownish stain. The wax, with the insects beneath, surrounds the twig as the wax 
does the wick of a candle; the whole mass is about 20 mm. diameter, that of the 
t vig being about 5 mm. 

“ Mr. King found the antennae to measure thus in 1 :— 


4. halide Abeta 4 diag 029 ml ¢) Haan ©: i (3) a) 
Pou ok er. GS 8! 60 a8 ag 49 
Rees ee ag 4 gh BB! SOR gd 


“Found by Mr. Fuller at Richmond, Natal. The nearest ally is an undescribed 
species from Paraguay, collected by Professor Bruner.” (Ckll. & King). 


This species is not represented in the collection. 


226. Ceroplastes combreti, sp. n. 

Test of 2 about 3 mm. long, broad and high, conical, with the anterior side a little 
more precipitous than the posterior, which is somewhat excavate above. There 
are no plaques, but the whole body of wax is arranged in + distinct columns, three 
on each side. The apex is blunt, bearing an opaque white ridge surrounded by six 
opaque white spots—the tops of the columns. Between the two most prominent 
lateral ridges are the distinct white stigmatic bands which extend from the base 
to the crown. The colour, when fresh, is bright rose-red with darker transverse 
marks. 

The aninnae are 8-jointed ; range in w: (1) 37-44, (2) 56-61, (3) 34-48, (4) 27, 
(5) 37-4 ', (6) 20-24, (7) 17-24, (8) 37-41. 

Leg I: coxa 75, femur + trochanter 180, tibia 129, tarsus 85, claw 20. 


28 CHAS. K. BRAIN. 


Anal plates + semicircular, about 160u long and 85u broad. The derm is thin 
and transparent, with numerous small, scattered, simple glands, from some of which 
short, tubular projections arise. Stigmatic clefts with large numbers of very 
small, roundly conical, thimble-shaped spines. 

The distinct, long, opaque white lines, the reddish colour, soft wax, etc., suggest 
quadrilineatus of Newstead, but this latter species has 6-joited antennae and is 
obviously quite distinct by other characters of the test. 

Habitat: On stems of Combretum sp., De Wildt, Pretoria District ; collected by 
Claude Fuller, July 1918. 
Collection No.: 317. 


227. Ceroplastes destructor, Newst. 

Ceroplastes ceriferus (Anderson) Newstead, Bull. Ent. Res. 1, pp. 66, 195, 1910. 

Ceroplastes destructor, Bull. Ent. Res. vii, p. 26, 1917. 

“ Female test. White, creamy white or dirty white; exceedingly soft and con- 
taining an excess of moisture. Form irregular, with large but ill-defined gibbose 
protuberances ; sides usually with two narrow opaque lines of secretion from 
the stigmatic clefts. No trace of lateral plates. Length, 4-8 mm. 

“Female, adult. More or less hemispherical, with the sides often slightly com- 
pressed ; caudal process long; integument castaneous and highly chitinised, 
smooth and shining, and without fovea or lateral tubercles. Antennae of six 
segments, the 3rd being as long as the 4th, 5th, and 6th together; the last three 
segments with stiff and bluntly pointed, spimose hairs. Legs small; hind femora 
very short and often distinctly incrassate ; hind tarsi equal in length to the tibiae, 
or sometimes slightly longer. Claw very short ; lower digitules very long and stout ; 
upper digitules normal. Stigmatic clefts well defined, but relatively small; stig- 
matic spines very small and pointed, bases not constricted; basal attachment 
(disc) very large. Caudal process (after maceration) transparent and somewhat 
flexible ; sides with an irregular double row of short spinose hairs, and in addition 
to these there are two pairs of longer hairs (one pair of which is twice the length 
of the others) slightly ventral to the row of short ones and towards the distal 
extremity. Anal lobes short and highly chitinised. Dorsal pores very small, 
rather widely separated. Ventral integument opposite the caudal process, with 
rather extensive groups of circular pores, many of which, in well cleared specimens, 
are linked together with lines of dark chitin. Length 4-7 mm.” (Newstead.) 

Habitat: On custard apple and avocado, Nelspruit, Transvaal; collected by 
D. Gunn, September 1915 (Coll. No. 93). On syringa (Melia azedarach), Bechuana- 
land, October 1918 (Coll. No. 336). 

Collection Nos. : 93 and 336. 


228. Ceroplastes egbarum, CkIl. 

Ceroplastes egbarum, Ckll. The Entom. xxxii, p. 127, 1899. 

Ceroplastes cristatus, Green, Ann. Mag. N.H. (7) iv, p. 190, 1899. 

Professor Cockerell’s description is as follows :— 

“Waxy female scales often crowded on the twigs, two or more coalescing ; 
about 11 mm. long, 10 broad, and 6 high, the wax extremely thick, not at all divided 
into plates, snow-white, here and there with a suffused pinkish stain. 


THE COCCIDAE OF SOUTH AFRICA. 29 


‘* ©. Denuded of wax 53-7 mm. long, 4 broad, 2}—3 high, very dark, with a dorsal 
hump but no lateral humps; anal horn a mere mammiform prominence. Boiled 
in caustic soda the denuded females give a purple colour, which on dilution with 
water appears pink, and soon forms a flocculent pink precipitate. On adding 
nitric acid a flocculent white precipitate appears, but the pink precipitate is not 
altered. Skin after boiling remains yellowish brown, chitinous, with scattered 
minute gland-dots. Stigmatic areas with numerous crowded gland-spots, and 
many short and rather thick simple spines, but no capitate spines. Legs dark 
brown, the parts measuring thus in w: Coxa, 120; femur with trochanter, 180: 
tibia, 128; tarsus with claw, 96 to 114. Tarsal digitules 60, slender, with a small 
knob. Claw digitules with very large round knobs, extending about 15u beyond 
tip of claw. Antennae apparently only 6-segmented, but the segmentation towards 
the end very obscure. The segments measure in w: (1) 45; (2) 60-69; (3) 66-78 ; 
(4) 51; (5) 69; (6) 72. Segment 5 has a deep notch which makes it look as if 
divided into two. 

“Young larvae under female about 4301 long and 230 broad, tinged with a warm 
reddish colour. Male scales small, elongate, and glassy.” 

In the description of C. africanum var. cristatus Green states that specimens from 
Natal differ from africanum (1.e., mimosae) only in the presence of a small dorsal crest 
corresponding to the position of the central scar. They were, however, larger, 
being 12 mm. indiameter. This would seem to indicate, moreover, that the antennae 
may be 6, 7, or 8—jointed. 

Habitat: On Acacia, Natal. 

Collection No.: 87. 


229. Geroplastes egbarum fulleri, T. & W. Ckll. 

Ceroplastes egbarum subsp. fullert, T. & W. Ckil., The Entom. xxxv, p. 113, 1902. 

Adult 9, with waxy covering about 14 mm. long, 12°5 mm. wide and 7 mm. high. 
Waxy covering regularly domed with a slight depression in centre. Margins widely 
crenulate, with usually two conical deflected waxy masses which clasp the stem. 
Stigmatic cleft waxy appendages conspicuous, snow-white, long and _ slender. 
There are two of these on each side, usually 3 to 4°5 mm. long, about 1 mm. thick 
at attachment to waxy covering, but gradually tapering to their extremities. They 
usually he closely pressed to the stem of the host plant. The colour of the waxy 
covering is whitish to coral, pink, with distinct brown patches on the intermediate 
area and lighter zones around the dome. 

Female, denuded of wax, 7 mm. long, 5°5 mm. broad and 4 mm. high. Colour 
coral pink, more yellowish than the darker coloured wax. The extreme frontal 
margin, the stigmatic clefts and the caudal projection are dark castaneous and 
shiny. In older specimens, after oviposition, the integument becomes brown. 

The venter is flat or concave with the median zone sunken and segmented. The 
4 white stigmatic bands extend inwards to this sunken area. The lateral margins 
are slightly excavate, the upper edge of the excavation being formed by seven 
depressed conical projections from the intermediate area. The anterior of these 
extends forward until nearly level with the front edge of the rounded anterior lobe. 


30 CHAS. K. BRAIN. 


The three lateral projections of each side are shorter and more bluntly pointed. 
The depressed area which separates the central cone from the intermediate area is 
coarsely punctate. The central cone is regular, elongate, oval at the base and 
roundly pointed at the apex. There is no sunken area in the centre. 


The caudal projection is exceptionally short, bluntly conical, projecting at an 
angle of about 45° with the ventral surface. 

Antennae 8-jointed; range in mw: (1) 44-51, (2) 48. (3) 58-68, (4) 34-37. (5) 
27-34, (6) 20-24, (7) 20-24, (8) 37-48. 

Integument clear, hyaline, except for the extreme margin, marginal expansions 
and caudal prominence, which are densely chitinised. The anterior margin is broadly 
rounded with a distinct parallel-sided chitinous band. Lateral margins with two 
almost circular expansions on each side. These are much larger than those found 
in mumosae. 

Habitat: On “ monkey rope,” Natal coast, Umbilo and Equeefa Rivers, 

Collection No.: 88a. 


230. Geroplastes eucleae, sp. n. (Plate u, fig. 239). 

Adult 9 tests sometimes single on stem, often aggregated in dense masses, Test 
of adult 2 about 6 mm. long, 5 mm. wide and 5°5 mm. high, without plaques but 
with the lower portion forming a wrinkled fold at the base of a highly conical dome. 
The colour is a delicate green, when alive, with the stigmatic bands conspicuous 
(fig. 239); when dry, it is semi-transparent, greenish yellow, with two white thin 
streaks on each side just above the stigmatic clefts. The central dome is pointed, 
without any central pit or depression, and is distinctly separated from the lower 
portion of the test by a groove. There is no indication of a caudal prominence on 
the test. 

Female, denuded of wax, smooth, regularly domed, without caudal prominence, 
chitin pale brown and moderately thin ; portion surrounding the anal plates deep 
castaneous. Caudal prominence rudimentary, indicated by deeper coloured 
chitin. When cleared, the derm is moderately chitmous. That of the dorsum 
is very finely rugose with very scattered small, transparent spots. The denser, 
marginal folds have in addition a few larger holes. 

Antennae 6-jointed; range in w: (1) 24-30, (2) 37-40, (3) 91, (4) 17-24, 
(5) 24-27, (6) 40. 

The legs are moderately developed, normal. 

Stigmatic clefts with a series of short, conical, thimble-shaped spines, which 
extend in a single row for some distance along the margin on either side of the cleft, 
where they compose a double row. About the middle of the group is a large spine, 
about twice as large as the others. Within the double row is a collection of small 
simple glands like the circumgenital glands of the D1asPINaz. 


Habitat: On stems of several native shrubs, including Huclea sp., Ochna sp. ?, 
Pavetta sp. ?, etc., Pretoria ; collected by Miss E. Impey, January 1915. 


Collection Nos. : 90 and 342. 


THE COCCIDAE OF SOUTH AFRICA, 3] 


231. Geroplastes longicauda, sp. n. 

Adult 2 covered with a very thick layer of soft, white wax forming a test like a 
large ceriferus specimen, i.e., a little more elevated than egbarum. Largest specimen 
seen measured 18 mm. long, 11 mm. wide and 12 mm. high; marginal area 
prominent, forming a wide fold at the base of the central dome. The waxy 
appendages from the stigmatic clefts only project slightly from the main mass 
of the fold. 


Adult 9, denuded of wax, bright brown in colour, about 5 mm. long without 
caudal process, which alone measures 3°5mm. The body is + star-shaped with 
three short, lateral pointed spurs and one anterior. These,are sharper and more 
prominent than those of ceriferus. The dorsum is very convex, rising with straight 
sides to an acute point. The caudal process is exceedingly long, two thirds the lengt' 
of body, piceous, shghtly tapering, and extends in a horizontal direction, 1.e., flat 
along the twig. 

Cleared and mounted, the insect is remarkable for the broad oval, thin, trans- 
parent body with a very long dense black tail. The integument is uniformly hyaline. 
without chitinous marginal discs such as those found in fulleri, etc. In stained 
material the integument of the dorsum illustrates a strange segmentation -+ in 
plates, 5 elongate transverse median ones and shorter laterals. 

The antennae are 7 or 8-jointed, e.g., (1) 40, (2) 37, (3) 40, (4) 47, (5) 68, (6) 30, 
(7) 27, (8) 374; or (1) 23, (2) 34, (3) 40, (4) 88 (with pseudarticulation), (5) 27, (6) 27, 
(7) 37. 

Stigmatic cleft thin, hyaline, with a patch of scattered, short, thimble-shaped 
spies and, within this, a large group of simple glands similar to the circumgenit.l 
glands of the DIASPINAE. 

Legs comparatively short, otherwise normal. 

Habitat: On stems of native shrub; collected by C. Fuller, Natal Coast, July 
1915. 

Collection No. : 334. 

This species is very similar in many respects to C. ceriferus but may be 
readily separated by the larger size of the adult 9 test, the comparatively longer 
caudal process and the 7 or 8-jointed antennae. C. ceriferus has antennae 5 or 6- 
jointed, usually 6. 


232. Ceroplastes mimosae, Sign. (Plate ii, fig. 241). 

Ceroplastes mimosae, Sign., Ann. Soc. Ent. Fr. (5) ii, p. 46, 1872. 

Ceroplastes africanus, Green, Ann. Mag. N. H. (7) iv, p. 188, 1899. 

“Tnsects crowded on the stems of the plant, so much so that the waxy covering 
of adjacent individuals becomes more or less confluent and the normal form of the 
test is difficult to determine. The tests appear as rounded masses of cream-coloured 
wax, each with a more or less distinct nipple-like prominence at the apex bearing 
a small spot of whiter substance. 

‘ The usual opaque white bands from the spiracular regions are present, but very 
inconspicuous, scarcely extending beyond the margin. In some specimens a series 


of CHAS. K. BRAIN. 


of impressed arches on the sides of the test marks the position of the marginal 
plates. The waxy coating being thinner on the impressed parts, the arches appear 
darker, the colour of the body of the insect showing through the covering-matter. 
An isolated test averages 7°75 mm. long, 6°50 mm. broad, 5°75 mm. high. 

“Female, denuded of wax, reddish brown to dark brown, the whole surface 
strongly chitinised ; irregularly globose ; apex often with an oblong scar correspond- 
ing with the position of the early larval pellicle, but which becomes almost 
obliterated in the oldest examples. In the early adult the median is separated 
from the marginal area by a more or less distinct furrow, which is particularly 
marked where it meets the anal tubercle. In the older examples only this hinder 
part of the furrow remains. Cephalic area constricted off from the globose body, 
forming a trowel-shaped projection in front. Spiracular clefts deeply indented, 
thickly set with small conical spines, not constricted at the base. Marginal hairs 
very small, few and inconspicuous. Anal scales minute, inner edge straight, base 
and outer edge together forming a semicircle. Anal tubercle blackish, directed 
upwards. Derm with numerous glandular pores, which are more distinct on the 
darker marginal area. Antennae with either 7 or 8-joints. It is difficult to say 
which is the normal number, as the two varieties are about equally represented 
in the series under examination. With the 8-jointed form the formula runs :—3, 
(1, 2), 8, 4, 5, (6, 7). When there are seven joints only the formula is 3, (1, 2, 4), 
7, (5, 6). In this latter case there is a tendency for the fourth joint to separate 
into two, and there is always a more or less distinct false joint in the terminal 
segment. Legs well developed; tarsus more than half length of tibia. Foot 
with 4 digitules, the unguals broadly spatulate, the tarsals fine knobbed hairs. 

“Length of fully developed female 5°50 mm., breadth 5:0 mm., height 4°25 mm. 

“The male insect is unknown in any stage.” (Green.) 

The material I have examined has a characteristic odour and has uniformly 
8-jointed antennae, with the following range in 4: (1) 27-34, (2) 41-48, (3) 58-68, 
(4) 27-41, (5) 17-87, (6) 17-24, (7) 20-24, (8) 34-37. 

Habitat : On Acacia karroo, Cape Colony ; common but local. 

Collection No.: 85. 


233. Ceroplastes myricae (Linn.). 

Coccus myricae, Linn., Syst. Nat. Ed. xu, i, p. 741, 1766. 

Columnea myricae, Targ., Catalogue, p. 35, 1869. 

Ceroplastes myricae, Sign., Ann. Soc. Ent. Fr. (5) 1, p. 39, 1872. 

“ Habitat ad Cap. B. Spei, in Myrica quercifolia. 

““Magnitudo pisi minoris, semi-ovatus secundum perpendiculum, pallide incarn- 
atus, vertice obtuse acuminatus cum poro tenuissimo, postice supra marginem 
etiam porus est, margo cartilagineus, crassior albus, utrinque circiter septem torutis 
protuberans. 

“Dans Olivier, Encyclopédie, VI, 96, 8, nous trouvons une description presque 
identique : la femelle est presque de la grandeur d’un petit pois, le corps est d’une 
couleur rouge pale et de forme demi-ovale, le vertex est elevé et percé d’un petit 
point, tout le bord est cartilagineux, épais, blanchAtre, marqué de chaque cété de 
petits cordons élevés. 


THE COCCIDAE OF SOUTH AFRICA, 33 


“Ce sont ces sept cordons élevés qui, spécifiant bien ’espéce, nous empéchent de 
Yattribuer aux nombreux individus que nous possédons et decrivons sous le nom 
de C. Vinsonu.” (Sign.) 

Myrica quercifolia, Linn. (Myricaceae) is a near relative of the waxberry 
plant which is common on the Cape Flats. Mr. C. W. Mally, the Entomologist 
for the Cape Province, has recently kindly examined a large number of plants in an 
endeavour to re-discover this species of Ceroplastes, but as yet without success. 


234. Ceroplastes pallidus, sp. n. 

Test of adult 2 to 85 mm. long, 6 mm. broad and 4:5 mm. high; very much 
like alarge C. rusci in form, with 8 lateral plaques and median dome. The lateral 
plaques, however, are without “ nuclei ” and are a little more perpendicular. In 
old specimens the waxy covering is pale, semi-transparent, yellowish, or having 
an indistinct greenish tint. The central dome is moderately elevated, somewhat 
tapering, slightly glossy, with longitudinal and concentric striae. The central 
“nucleus” is elongate, glossy and a little darker in colour. Stigmatic clefts 
indicated by small white dots. 

When cleared and mounted the integument is all thin and hyaline, without anterior 
or marginal thickenings and with only a small area surrounding the anal lobes chiti- 
nised. 

Stigmatic clefts shallow, not chitinised, with short, conical, pointed spines which 
are in a single row for a short distance on each side, but broaden out to form a small 
triangular patch opposite the spiracle ; at no point, however, are there more than 
5 rows. Within this series is a compact group of simple glands like the circum- 
genital glands of the Drasprnaz and also a few smaller glands and short simple hairs. 
The remainder of the margin has a scattered, single row of short, slender, sharply 
pointed spines. 

The antennae are 8-jointed; range in w: (1) 37-44, (2) 37-47, (3) 44-52, (4) 
34-44, (5) 51-56, (6) 27-32, (7) 27-34, (8) 35-47. 

Legs well developed, normal; tibia long, with a constriction at about the middle. 

The caudal tubercle is comparatively small, a mere plate with the chitin appearing 
-- streaky and somewhat perforate. 

Habitat : On fig, Church Square, Pretoria. 

Collection No.: 102. 


235. Geroplastes quadrilineatus, Newst., var. simplex, nov. 
Adult °° clustered in large masses on the twigs of the host plant. @ test about 
5 mm. long, 4°5 mm. broad and 3 mm. high, very dark in colour, almost black, with 
the thinner parts of the wax appearing reddish. Marginal fold corrugated, lighter 
and more transparent in colour than the central dome, with two very distinct lines 
of opaque white on each side. The central portion is comparatively flat and trun- 
cate, separated from the marginal fold by a deep furrow in dry material, with the 
apex hollowed into a shallow cup containing an opaque white ridge. The ridge 
of this cup-like depression is not regular but consists of an anterior transverse ridge 
and two posterior lateral spurs, and the floor usually slopes gently forward. The 
(681) , C 


ot CHAS. K. BRAIN. 


anal opening to the test is surrounded by a distinct circular ring which is itself 
sunken in a deep depression with broadly rounded sides. In dry specimens the whole 
waxy material is roughened, hard and very brittle. The pair of larger divergent 
pytiform bodies referred to in the description of quadrilineatus are not present. 

When cleared of the waxy covering the insect is of a characteristic form, with the 
dorsum uniformly rounded and smooth and the marginal area produced in a series 
of broadly rounded bead-like lobes. The caudal prominence is extremely rudimen- 
tary, appearing as a flatly rounded hump, black in contrast with the deep red 
brown of the remainder of the insect. The integument, when cleared, is dense 
and is pitted like that of many species of Sazssetia. The stigmatic spines are in 
an elongate row around the shallow cleft, the series becoming double in the centre. 

The antennae are 6-jomted, three being very long and variable, e.g. :—(1) 27, 
(2) 37-40, (3) 88-119, (4) 20, (5) 27, (6) 44-51 u. 

Legs long, normal. Chitin of caudal protuberance appears as though bossed 
w.th a honey-comb pattern. 

Habitat: On stems of Rhus sp. (probably R. vimanalis), Victoria West, C.P.; 
collected by Mr. van Heerden, October 1915. 

Collection No.: 346. 


236. Ceroplastes rusci (Linn.) 

Coccus ruscir, Linn., Syst. Nat. Ed. x, 1, p. 456, 1758. 

Coccus caricae, Bern., Mem. Acad. Marseille, p. 89, 1773. 

Coccus artemisiae, Rossi, Mant. Ins. 11, pp. 56, 514, 1794. 

Calypticus radvatus, Costa, Faun. Reg. Nap. Coce., p. 12, 1835. 

Calypticus testudineus, Costa, Faun. Reg. Nap. Cocc., p. 12, 1835. 

Calypticus hydatis, Costa, Faun. Reg. Nap. Cocc., p. 14, 1835. 

Columnea testudinata, Targ., Atti dei Georgofili, n.s. xi, p. 31, 1866. 

Coccus hydatis, Targ., Studi sul. Coce., p. 12, 1857. 

Columnea testudinforms, Targ., Studii sul. Cocc., pp. 8, 11, 12, 1867. 

Chermes caricae, Bdv., Ent. Hort., p. 320, 1867. 

Ceroplastes rusci, Sign., Ann. Soc. Ent. Fr. (5) u, p. 35, 1872. 

Lecamum artemisiae, Sign., Ann. Soc. Ent. Fr. (5) i, p. 37, 1872. 

Adult 9. Test about 7-5 mm. long, 5:2 mm. broad and 5 mm. high, regularly 
domed, rounded in front, rather excavate behind. Colour greasy white, suffused 
with purplish red. Dorsal dome and margins of plaques lined with greenish grey 
lines ; anal pore and middles of plaques of same colour. Stigmatic bands, 2 on each 
side, pure white, broad at stem and tapering inwards. Middle of dorsum with a 
depression in which there is an elongate, glossy white prominence similar to those 
in centres of plaques, but longer. 

Antennae 6-jomted; range in uw: (1) 34-40, (2) 37-40, (3) 114-120, (4) 17-24, 
(5) 20-27, (6) 44-57. 

Leg II: coxa 102; femur + trochanter 170; tibia 120; tarsus 85; claw 20wu. 
Margin of sides of body with a single row of small thimble-like spines ; amongst 
these are mixed a few simple spines. Stigmatic clefts very shallow, not chitinised, 


THE COCCIDAE CF SOUTH AFRICA. 35 


with a few additional spines like those of the marginal series and a small group 
of simple glands like the circumgenital glands of the Diaspinag. Anal plate heavily 
chitinised, with scattered perforations. 

Halitat: On stems of quince, Stellenbosch, C.P.; collected by F. W. Pettey, 
July 1916. 

Collection No.: 91. 


237. Ceroplastes tachardiaformis, sp. n. 

Adult tests aggregated in huge masses on stems of the host-plant, often com- 
pletely covering the stems for a distance of several inches. 

© test Tachardia-like, globular, slightly flattened above, hard, thin, brittle, 
almost transparent resinous brown, due to the colour of the insect within ;_ without 
protuberances but with a slight apical depression containing the opaque white 
larval exuvia. 

Adult 2, denuded of wax, moderately dense, globular, smooth and shiny. Caudal 
protuberance short, very dense, surrounded by a coarsely perforated plate. 

Antennae 6-jointed, e.g. (1) 34, (2) 27, (3) 74, (4) 20, (5) 17, (6) 30u. 

Legs short, e.g., coxa 50, femur ++ trochanter 110, tibia 78, tarsus 50. 

Integument moderately dense, appearing, under the high power, finely rugose, 
with numerous, widely scattered, small, transparent pores. Stigmatic clefts almost 
obsolete, indicated by a small group (8 or 10) of short obconical spines and a few 
simple glands. 


Habitat: On rhenosterbosch (Elytropappus rhinocerotis, Less.) ; collected by 
Messrs. Watermeyer Bros., Aberdeen, C.P., November 1915. 


Collection No.: 94. 


238. Ceroplastes zonatus, Newst. 

Ceroplastes zonatus, Newstead, Bull. Ent. Res. vii, p. 32, 1917. 

“Female test. Broadly ovate in outline, highly convex; marginal plates 
very faintly indicated, but apparently without nuclear spots; dorsal plate very 
large, with a central nuclear spot of white wax ; cephalic margin slightly clypeate ; 
lateral margins in very old examples with a pronounced foot-like extension from 
each of the stigmata, from which there extends a thick white waxen appendage. 
In the younger forms the foot-like extension is wanting, but the white waxen appen- 
dages are present and always porrected. Colour creamy white, suffused with very 
pale brown; dorsal plate surrounded by a shaded wavy zone of dark brown and 
brownish black, with here and there a suffused patch of dull flesh-colour. In very 
old examples the zone of colour extends to the margins and is of a shining madder- 
brown to piceous colour. On the removal of the outer surface of the test with 
chloroform, it is seen to be divided into seven areas by pale orange-coloured lines ; 
a central polygonal area, corresponding to the area occupied by the dorsal plate, 
from the angles of which radiate to the margin single lines marking off the areas 
of the lateral and cephalic plates. 

“Female adult (denuded of the test). Ovate; cephalic margin clypeate ; 
dorsum low and wrinkled; two large, bilateral, submarginal extensions, both 
longitudinally striated : the space between these extensions of the body-wall and 

(681) eo 


36 CHAS. K. BRAIN, 


the margin markedly constricted. Dorsum with a large keel-like process. Caudal 
process very short and conical. Stigmatic clefts deep. Antennae of eight seg- 
ments ; the articulations relatively very broad ; 3rd about equal in length to the 
7th and 8th together ; a very long hair on the 2nd and 5th, and a slightly shorter 
one on the 8th ; there are two spines on the 8th and one on the 7th. Legs normal. 
Stigmatic spines covering a large and somewhat pyriform area, the length of which 
is nearly equal to twice the length of the antennae ; the spines, with the exception 
of a small group near the stigmata, are obconical and the space between them with 
bands of dark granular bodies, which collectively form a polygonal reticulation ; 
the small proximal group of spmes are longer than the others and pointed. No 
trace of marginal spines. Derm thin and transparent after maceration ; rather 
thickly set with minute pores and minute scattered spines. Caudal process sur- 
rounded by a porose zone of brown chitin. Length 3°9-4°6 mm.” (Newstead). 

The above description does not refer to the mature female forms which attain, 
with the waxy test, 14 mm. long, 12 mm. broad and 8 mm. high. The colours 
remain the same except that the lighter parts become yellowish. 

When mature the denuded female is densely chitinous, brownish black, with the 
dorsum smooth, shining. In boiling KOH it stains the liquid deep purplish 
brown. The antennae are sometimes 7-jomted with joint 4 very long, obviously 
4 + 5 of the 8-jointed form. 

Halitat: On Acacia sp., Pretoria, November 1914. 


Collection No.: 344. 


Genus Inglisia, Mask. 
Adult female more or less conical, covered above by a glassy shield which is 
divided into plates and striated with rows of air cells. Legs and antennae well 


developed. 


239. Inglisia elytropappi, sp. n. 

Test of adult 9 small, 1°6-2 mm. long, 1°2 mm. wide and high, like a small bivalve 
shell with its hinge uppermost standing on the stem. It is very like a small zizypha 
test but paler in colour, with the upper angles of the two halves smooth instead of 
tuberculate. The colour is white to pale buff, shiny, often pearly, with the vertical 
striae conspicuous. 

2, with the test removed, shiny dark brown, of the same general shape as the test, 
with the apex depressed between two rounded lateral humps. 

Cleared and mounted the body is hyaline. The margin has a close-set row of 
short conical spines with broad bases and moderately sharp points. The stigmatic 
clefts are obsolete, but their presence is indicated in the marginal row of spines by 
the addition of a single, slightly longer spine with a narrower base. 

The antennae are rudimentary, appearing in varied forms with very indistinct 
segmentation ranging from 4 to 7-jointed. When 7-jointed the segments are 
generally very short—mere rings. The measurement of such an antenna gave 
the following in wu :—(1) 17, (2) 6, (3) 17, (4) 6, (5) 10, (6) 6, (7) 6. 


THE COCCIDAE OF SOUTH AFRICA. 37 


Legs rudimentary, appearing as though composed of three, almost equal, cylindri- 
cal segments with a minute claw. The total length varies between 70 and 90. 

Habitat: On the thinnest twigs of rhenosterbosch (Hlytropappus rhinocerotis, 
Less.), Groot Drakenstein, Somerset West and Cape Flats, C.P. (Cape Coll. No. 
1244). The stems of the host-plant are thickly covered with “ sooty”’ fungus, 
apparently grown on secretion from the numerous specimens present. 


Collection No.: 100. 


240. Inglisia geranii, sp. n. (Plate iv, fig. 249). 

Insects congregated on the main stems at or near the nodes. Adult insect, 
with test, about 2°5 mm. long and 1°3 mm. broad at the base, brown, with the air 
spaces of the test almost colourless and appearing as transverse lines radiating from 
the centre of each half of the test. The test is composed of two similar halves, each 
of which simulates a shell or tortoise-shell, with their apices widely separated. The 
median line at the point of union of the two is almost flat, very little depressed. 
The test easily flakes away from old specimens and is then white, almost hyaline. 

The antennae are 7 or 8-jointed ; range in « :—7-jointed: (1) 24-31, (2) 14-20, 
(3) 41-48, (4) 17-20, (5) 14-20, (6) 14-17, (7) 20-24; 8-jointed: (1) 24, (2) 17, (3) 
31-34, (4) 14, (5) 14-17, (6) 14-17, (7) 17, (8) 24. 

Leg I: coxa 68; femur + trochanter 136; tibia 109; tarsus 85u. The tro- 
chanter has a long (90m) spine. The upper digitules are long and slender, hardly 
perceptibly clubbed ; lower digitules comparatively short and slender. 

Anal plate about 115 long, with 2 or 3 stout spines. Margin with a single row 
of sharp pointed spines of varying lengths ; these are all broad at the base and taper 
rapidly to the point. Stigmatic cleft with a single spine a little longer than the 
longest of the marginal series and a little less tapering. 


Habitat: On geranium, King Williamstown, C.P.; collected by A. Kelly, March 
1916. Fresh material sent by Mr. J. Hobson, King Williamstown, May 1916. 

Collection No.: 99. 

This species is remarkably close to Inglisia theobromae, Newst., which 
_ was described on cacao from Uganda, but it is smaller, and joint 3 of the antennae 
is apparently always longer than 4. It is also somewhat like J. bivalvata, Green, 
but the dorsal shield, formed by the inner sides of the two halves of the test, is wider 
and more oval. 


241. Inglisia zizyphi, sp. n. 

Test of adult 92 shaped like a small bivalve shell, standing erect, with the two 
halves separated by a conspicuous furrow, and the hinge uppermost, represented 
by two small, rugose “crowns” of the two halves. The lower margins, which 
rest on the stem, are surrounded by a slight fringe of fine glassy filaments. The 
test is pale horn-colour, with the distinct vertical striae appearing slightly iridescent. 
Length 2 to 3 mm.; width 15 to 2 mm.; height about 2 mm. 

Adult 9, with the waxy test flaked off, red-brown, of the same shape as the test 


but with the dorsum shiny and wrinkled. 


38 CHAS. K. BRAIN. 


Cleared and mounted the integument is thin and hyaline. Margin with a dense 
series of stout conical spines, so closely set that the row appears, in places, double. 
The derm is clear except for a double row of large simple glands extending along the 
median line from the anal plates to the middle of the dorsum. 

Antennae 5 or 6-jointed, the individual segments badly defined even in stained 
preparations ; range in «:—0d-jointed : (1) 24, (2) 14, (3) 41-51, (4) 24-34, (5) 24-27; 
6-jointed : (1) 20-24, (2) 10-14, (3) 48-51, (4) 10-17, (5) 14, (6) 20-24. 

Legs short but normal, e.g., coxa 41; femur + trochanter 102; tibia 75; tarsus 
58; claw 17u. Anal plates about 90u long, each with two fairly stout spines about 
40u long. Stigmatic clefts obsolete. 

Larva about 390u long; antennae 6-jointed ; caudal setae very long (170,). 

Habitat : On Zizyphus sp., Pretoria ; collected by the writer, December 1914. 

Collection No.: 101. 

This species is very close to I. conchiformis, Newstead, but is smaller and has 
an antennae of 5 or 6 segments instead of 7. I thought at first that I was dealing 
with young individuals of Newstead’s species but mounted specimens contain well 
developed embryonic larvae. 


. 


Genus Cryptinglisia, Ckll. 


“* A Lecaniine Coccid having a glassy covering containing air-spaces, and retaining 
the legs and antennae (7 or 8 joints) in the adult. Living in galls on the roots of 
Vitis. Differs from Inglisia in its mode of life; in the glassy scale not being divided, 
tortoise-like, into plates ; and in the air-cells running together, forming long air- 
spaces. Larva ordinary, with six large bristles on the cephalic margin. Male 
unknown.” (CkIll.) 


242. Cryptinglisia lounsburyi, Ckll. 

Cryptinglisia lounsburyt, Ckll., The Entom. xxxiil, p. 173, 1900. , 

Cryptinglisia lounsburyi, Lounsb., Rep. Ent. Cape Good Hope, p. 54, 1900. 

“ Adult female about 24 mm. long, soft, shiny, very dark brown, covered with a 
semi-transparent, brittle, glassy scale. Skin transparent and colourless on boiling 
in KOH; mouth-parts moderate, rostral loop not very long ; margin with a row of 
simple spines, brownish, about 24, long, placed close together ; anal lobes ordinary, 
about 160u long, yellowish brown, surrounded basally by a large, thick, dark brown, 
chitinous plate, more or less semilunar in form, with the ends produced ; a row of 
small round glands in the middle line from one end of the body to the other, but best 
developed posteriorly ; antennae and legs pale ; legs ordinary, femur plus trochanter 
about 120, tibia about 96, tarsus about 78, claw about 20u ; claw-digitules about as 
long as claw, with large knobs ; tarsal digitules long, with distinct knobs. Antennae 
7 or 8-jointed, having three types, thus: (1) 7-jointed with a short 3, all the joints 
subequal, 21 to 30u; (2) 7-jointed with a long 3, which is about 41 long; (3) 8- 
jointed, with 2 quite short, and 3 and 4 each about 30u long. The terminal joint is 
always short, 21 to 26. 


THE COCCIDAE OF SOUTH AFRICA. 39 


“These insects occur underground on the roots of grape vines, living in galls 
which are more or less globular, 4 to 5 mm. diameter, dark, rough and often nodulose 
on the outside, often aggregated together in numbers, or even coalescing, so that the 
root presents a nodulose thickening 6 or 7 mm. in diameter and over 20 mm. long. 
On breaking open the galls, which are quite hard, one finds a cavity containing the 
Coccid. Small stones are frequently embedded in the sides of the galls. 

“ Hab. Constantia, Cape Colony, at the roots of Stein and Reisling grapes (Vitis 
vinifera). Mr. Chas. P. Lounsbury, sending the specimens, says; ‘ None were 
observed more than eight or nine inches from the surface, and all were on fibrous roots. 
As you will observe from the specimens, they are somewhat gregarious ; ofttimes. 
one or two rootlets will be quite covered, while all the others are free. Most of the 
infested vines were backward in growth—some almost dead ;_ but their condition, 
I think, is due to other causes than the attack of the insect. Some apparently 
healthy vines were noticed to be affected.” (CkIl.) 

This species has not been re-discovered. 

Collection No.: 105. 


Genus Parafairmairea, Ckll. 
Female scale divided by a longitudinal, median suture into two halves, each with 


minute grooves radiating from its apex but not striated with air cells. Legs and 
antennae well developed, the latter 7 or 8-jointed. 


243. Parafairmairea patellaeformis, sp.n. (Plate iv, fig. 250). 

Adult ° dull brown, 9 mm. long and 5 mm. broad, covered above by a stout shield 
which is divided longitudinally into two halves. This shield has the wavy lnes and 
exact appearance of one of the common shells of the more elevated “ Patella ” type. 

With the covering removed the insect is glossy brown with wrinkles and ridges 
radiating from a two-fold dorsal peak to the margin. 

Antennae 7 or 8-jointed ; range in u :— 

i, II. lg bal ts va Ve. Oe eee 
20-31 20-24 41-68 34-41 20-27 14-24 20-34 - 
20-31 20-24 44 #17-34 27-34 17-20 1420 24-34 

Leg 1: coxa 88-102 ; femur + trochanter 187-238 ; tibia 160; tarsus 90; claw 
approximately 20. 

Anal plate about 1604 long. The integument is thin and hyaline, without con- 
spicuous hairs or glands. The margin has a single row of short, conical spines of the 
Inglisia ty pe, amongst which are intermingled, at intervals, smaller spines of a thinner 
type. The stigmatic clefts are obsolete, merely indicated by a scant series of small 
circular, simple glands extending inwards from the margin towards the spiracle. 

Remarks : This is the third Parafairmairea to be described. The other two were 
described on grass, one from France and the other from Surrey, England. 

Habitat: On stems of Acacia karroo; collected by A. E. Kelly at Port Alired, 
C. P., March 1915. 

Collection No.: 98. 


40) CHAS. K. BRAIN. 


Genus Geroplastodes, Ckll. 
Female scale convex, but not cone-shaped ; not divided into two halves, nor into 
distinct plates, but rough or beset with protuberances. © with antennae 7- or 
8-jointed. 


244. Ceroplastodes bituberculatus, sp.n. (Plate iv, fig. 251). 

Test of adult 9 about 4°5 mm. long and 3 mm. broad and high, white, with two 
prominent humps, one at each end of the median ridge; margin with a distinct 
white fringe. The test indicates the anal cleft and has a small prominence over the 
anal lobes. 

. Male scale dull white, with the median area denser, about 1°5 mm. long, divided 
into plates in the normal manner. 

Adult 2 with the waxy covering removed, dark brown, of similar shape to the test. 
Integument, when mounted, clear, hyaline, without glands or hairs. Margin with 
a single, close-set row of tubular spines somewhat of the Inglisia type but more linear 
and truncate. Stigmatic clefts obsolete but indicated in the marginal series of spines 
by the addition of three stigmatic spines ; laterals slender, a lictle longer than the 
marginal spines ; median stout, two and a quarter times as long as laterals (1161). 

Legs and antennae long, well developed. Antennae 8-jointed; range in yw: 
(1) 27-34, (2) 20-34, (3) 63-68, (4) 31-34, (5) 37-48, (6) 17-20, (7) 17-20, (8) 34-41 
(from 5 antennae). 

Ley I: coxa 78; femur + trochanter 176; tibia 120; tarsus 90; claw 24y. 
Tarsal digitules very long, slender, clubbed ; claw-digitules comparatively slender. 

Anal plates about 145 long, apex with 4 very stout blunt spurs. 

Habitat: On stems of native shrub, Somerset West, C.P.; collected by T. F. 
Dreyer, November 1906. Also at Stellenbosch (Fuller). 

Collection No. : 327. 


: Genus Idiosaissetia, nov. 
Adult 9 secreting a thin brittle covering of waxy material not divided into plates 
or two halves, and without air cells. Legs and antennae present but rudimentary. 
Anal cleft not median but to one side. 


Type, L. peringueyt, sp. n. 


245. Idiosaissetia peringueyi, sp. n. 

Adult 9 about 2°6 mm. long and 1°3 mm. broad, elongate, very convex, with 
almost perpendicular sides and sloping abruptly in front and behind. 

Female raised on a hollow waxy sheath which extends over the sides of the body, 
leaving the dorsum naked. The material is now ten years old and appears as though 
the waxy covering had once extended over the dorsum, but had become detached 
from it. If this were covered the insect would look like an Inglisia, except that 
the waxy covering is thin, pale buff-coloured and solid, 1.e., without air-tubes. 

Female, denuded of wax, densely chitinised, brown, shiny. 

When cleared and mounted the integument of the dorsum is moderately dense, 
with numerous + circular clear spaces as in Savssetia bitt of greatly varying size. 
The venter is thin and clear. The antennae are rudimentary, usually with about 


THE COCCIDAE OF SOUTH AFRICA. 41 


three indistinct joints, always exhibiting pseudarticulations. In the longest 
antenna seen, after staining, one could observe 6 ill-defined joints measuring 
(1) 17, (2) 14, (3) 14, (4) 14, (5) 14,(6) 20n. 

Leg I: coxa 34; femur + trochanter 85; tibia 85; tarsus 27; claw 17w. 

The margin has a scant series of short, curved, moderately stout spines, except 
near the anal cleft, where the spines are closer set, long (41), straight and very 
acute. The anal cleft is invariably thrown to one side and is not in the median 
line as in Lecanium. The anal ring has numerous (12?) hairs; anal plates about 
120ulong. Embryo-larva large, about 420, long, with 6-jointed antennae. Caudal 
extremity produced, with prominent caudal lobes, each with one long, stout seta 
(136) and several shorter spines. 

Habitat: On grass or thin reed. Label in tube reads: “ From Dr. Peringuey, 
S.A. Museum, Nov. 1908.” 

Collection No.: 140. 


Genus Membranaria, nov. 
Lecaniid, somewhat like Pulvinaria but with the cottony ovisac replaced by a 
membranous receptacle. Antennae and legs well developed, the former 7 or 8-jointed. 


Type, M. pretoriae, sp. n. 


246. Membranaria pretoriae, sp. n. (Plate iv, fig. 247). 

Adult 2 with ovi-receptacle about 5°5 mm. long, 2 mm. broad and 3 mm. high. 
Receptacle membranous, secreted from the margins, particularly the posterior 
margin, of the insect, so that, when completed, the insect, except the head end, 
is raised from the stem. The membrane consists of two coats, both thin, but 
which may be readily separated. The outer coat is pale, honey-comb yellow, not 
striated, and is produced from the upper margin of the insect. The inner coat is 
paler in colour and has longitudinal striae. The completed receptacle is hoof- 
shaped (fig. 247) with a distinct, dense, longitudinal, median keel. Its greatest 
length, along this keel, is about 3 mm. 

The adult ° is deep caramel-brown, sometimes speckled with black, glossy, with 
a rounded, median ridge and marginal corrugations and depressions. Cleared and 
mounted the adult 2 is moderately dense. Margin with a single row of simple, 
hair-like spines set wide apart. Submarginal area, except of anterior end, with a 
broad band of very numerous, small, gland pores, each with a curved, linear gland 
tube. Stigmatic cleft obsolete, but indicated by a pair of curious, short, broad, 
cup-like; protuberances. Anal ring with 8 hairs; anal plates surrounded in 
front by a dense, rugose, chitinous plate. 

Antennae 7 or 8-jointed ; range in mu :— 

: I III IV V VI VII Vill 
20-31 31-34 17-41 20-48 17-24 17-24 =. 24-27 — 
24-27 «27-37 = «24-3400 sd14-24 020-27 17-20 =16-20 24-27 

Leg I: coxa 102, femur + trochanter 238, tibia 153, tarsus 51, claw 20. 

Halntat: On crowns of grass, in front of Union Buildings, Pretoria ; collected 
by the writer, October 1914. 

Collection No.: 83. 


a9 % . 
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EXPLANATION OF PLATE I. 


Fig. 233. Lecanium elongatum, Sign., adult 929. 
234. Saissetia persimilis, Newst., adult 99. 


235. Pulvinaria mesembryanthemi (Vall.) Sign., adult 99 
with ovisacs. 


236. Filippia carissae, sp. n., adult 2 with ovisacs com- 
pleted and one ¢ puparium. 


237. Sarssetia subpatelliforme, Newst., adult 29. 


238. Hemilecanium theobromae, Newst., adult 99. 


PLATE I, 


PART 1. 


Yoru. AL 


BuLL. ENT. RESEARCH. 


SOUTH AFRICAN COCCID&~. 


EXPLANATION OF PLATE II. 


Fig. 239. Ceroplastes eucleae, sp. n., adult 9 9. 
240. Ceroplastes sp. 


241. Ceroplastes mimosae, Sign., adult 2 9. 


242. Pulvinarta aristolochiae, Newst., g tests and © 
ovisacs with larvae. 


243. Sarssetia sp., to illustrate dermal cells. 


244. Protopuluinaria piriformis (CkIL.) Lefroy, anal lobes of 
adult 9. 


BULL. ENT.- RESEARCH. Vo... XI. Part .1. Pcvate II, 


243. 


SOUTH AFRICAN COCCID&. 


a 
44% 


: 


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Vs ! \ ee , 
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iy yae - Ld 
aa 2 Sh 


EXPLANATION OF PLATE III. 


Fig. 245. Conofilippra subterranea, sp.n., Q ovisacs on 
roots ; a, 2 with top of ovisac tilted to one side ; 
b, section of ovisac with 9 ; c, swollen tibio-tarsa 
joint; d, antenna; e, stigmatic spines and 
glands ; f, anal lobes of 9. 


ET}. 


PLATE 


PART 1. 


VOL, (20n. 


Butt. ENT. RESEARCH. 


SOUTH AFRICAN COCCID/4 


EXPLANATION OF PLATE IV. 


. Allopulvinarva subterranea, sp.n., adult 29. 


. Membranaria pretoriae, sp. n., adult 2 9. 


. Protopulvinaria pirifornis (CkIl.) Lefroy, adult 
22. 
. Inglsia gerann, sp.n., young and adult 99. 


. Parafairmairea patellaeforms, sp.n., adult 99. 


. Ceroplastodes bituberculatus, sp. n., adult 99 and 


dS puparia. 


BuLt, ENT. RESEARCH. VOL. XI. PART 1. PLATE IY. 


SOUTH AFRICAN COCCID&. 


ae At Me ‘ ‘ ; ‘ ay, ; . suit, Om 6 : ¥ 
é * i \ ‘ whe f; ** J ot 4 ee SOS RL ea ee tee, P 
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Py 1 P ] ao ; = a" 
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43 


THE PERCY SLADEN TRUST EXPEDITION TO THE INDIAN OCEAN 
IN 1905, AND 1907-1909, UNDER Mr. J. STANLEY GARDINER, M.A. 


DIPTERA: TABANIDAE. 
By Major E. E. Austen, D.S.O. 


The material belonging to this family brought back by the expedition includes 
representatives of only four species, one of which, however, is apparently new. 
That three out of the four species obtained should belong to the Aegophagamyra- 
Bouverella group is not surprising, in view of the predominance of this group in 
the Malagasy Subregion. 

Through the courtesy of the authorities of the Percy Sladen Trust, the type of 
the new species described below has been placed in the British Museum (Natural 
History). 


PANGONIINAE. 
Genus Aegophagamyia, Austen. 
Ann. Mag. Nat. Hist. (8) 1x, p. 12 (1912). 


Aegophagamyia terticeps, Austen. 

P. ct., p. 15. 

Three 9 9, from Astove I., 1907 ; four 99 from Aldabra, 1908-9 (J. C. F. Fryer). 

The type and para-type of this species, in the British Museum (Natural History), 
are from Astove I. (P. R. Dupont). 

In the female sex, at any rate, this species bears a superficial resemblance to 
Tabanus albipectus, Big., for which on casual inspection it might easily be mistaken. 
The four specimens noted above as having been taken in Aldabra bear the same 
number as that attached to the examples of Tabanus albipectus collected in the 
same island, by Mr. J. C. F. Fryer, showing that the field note printed below under 
the heading 7. albipectus must be taken as applying to the present species also. 


Genus Bouvierella, Surcouf. 
Bull. Mus. Hist. Nat., Paris, 1909, p. 176. 


Bouvierella alluaudi, Giglio-Tos. 

Scione allawudi, Gigho-Tos, Ann. Soc. Ent. France, Ixiv, p. 357 (1895). 

Of this species, originally described from specimens from the Seychelles Is., the 
present collection includes no fewer than twenty-two examples, as follows :— 
One 3, one &, from Mahé, Seychelles, v-xu1. 1905 (J. S. Gardiner); eleven $6, 
five 99, from Mahé, Seychelles (5 gg, 2 29, from the Forét Noire district, x-xi. 
1908, and 6 93,3 929, from Cascade Estate, at 800 to 1,000 feet or more above sea- 
level, 1908-9); one 9, from Félicité, Seychelles, xi. 1908; 2 gg, from Praslin, 
Seychelles, x1. 1908; and one 9 from Cosmoledo Is., 1907 (H. P. T.). 

(681) D2 


44 FE. E. AUSTEN, 


Bouvierella inornata, sp. n. 

2.—Length (2 specimens) 10 to 11°2 mm. ; width of head 3°25 to 3°6 mm. ; width 
of front at vertex 0°4 mm., at lower extremity 0°25 mm. ; length of wing 9 to 10 mm. 

Sombrely coloured species, entirely devoid of markings ; ground-colour of dorsum 
of thorax drab* or dusky drab, that of dorsum of abdomen greyish fuscous ; front in 
© narrow, slightly narrower at lower extremity than at vertex, about six tumes as long 
as its width at upper end ; wings, especially portion beyond basal and anal cells, strongly 
suffused with brownish, stigma well marked ; coxae drab, legs otherwise blackish brown, 
extreme tips of femora, at least of those of front parr, ochraceous buff. 

Head: front, subcallus, face, sides of face and jowls drab, occiput smoke-grey ; 
vertex clothed with minute, erect, blackish-brown hair, sides of face with ochraceous- 
brownish hair, basi-occipital region with longer fine brownish hair ; frontal callus 
represented by a fine, dark, median line, commencing at some distance above level 
of lower inner angle of eyes, and not reaching anterior ocellus ; proboscis, including 
labella, narrow and elongate, total length of proboscis about two-thirds of that of 
head ; proximal joint of palpr: dark brown or brownish, terminal joint similarly 
coloured on outer surface, narrow, curved, elongate and very slender; antennae 
dark brown, first and second joints clothed above and below with short, blackish 
hair, expanded portion of third joint ovate or elongate ovate when viewed in profile, 
without any angle on upper margin, annulate portion of third joint narrow, curved 
and elongate. Thorax: dorsum clothed with short, recumbent brownish hair, 
which in front of prescutellar groove may be mixed with paler hair, scutellum bearing 
jonger brownish hair, or a mixture of brownish and ochraceous hair; pleurae and 
pectus agreeing in coloration with dorsum, and clothed with fine, brownish hair. 
Abdomen : dorsum clothed with short, appressed dark brown hair, venter agreeing 
with dorsum in coloration and hairy covering. Wings: lengths (relative as well 
as actual) of stalks of first and fourth posterior cells variable in different specimens ; 
stigma elongate, dark brown. Squamae light drab or drab-grey, borders drab. 
Halteres: knobs clove-brown or blackish brown, stalks paler (cinnamon-buff). 
Legs sparsely clothed with short or minute dark brown or blackish hair. 

Seychelles Is. Type from Silhouette (plateau of Mare aux Cochons, about 1,000 
feet, x. 1908) ; a paratype from Mahé (Cascade Estate, 800-1,000 feet), 1908-9. 

Bouvierella wmornata is distinguishable from all previously described species 
belonging to this genus, inter alia, by its uniformly sombre coloration, the ground- 
colour of the thorax and abdomen not being relieved by either lighter or darker 
markings. In particular, it is distinguished from B. brunnea, Surcouf (Bull. Mus. 
Hist. Nat., 1909, no. 4, p. 179), a species found in Madagascar, by its smaller size, 
and by the more uniformly dusky coloration of the body and legs, the dorsum of 
the thorax not being longitudinally striped with grey, and the legs not being reddish 
yellow, clothed with golden pile. 

In the length and slenderness of the proboscis, including the labella, the species 
just described agrees rather with Aegophagamyia, Austen, with which it constitutes 
in this respect an annectent link; as regards antennal characters and the shape 
of the first posterior cell, however, the species seems more correctly placed under 
Surcouf’s genus. 


* For names and illustrations of colours, see Ridgway, ‘‘ Color Standards and Color 
Nomenclature’ (Washington, D. C.: published by the Author, 1912). 


DIPTERA: TABANIDAE. 45 


TABANINAE. 
Genus Tabanus, Linn. 
Fauna Suecica, Ed. 1, p. 462 (1761). 
Tabanus albipectus, Bigot. 

T. albipectus, Bigot, Ann. Soc. Ent. France, (3) vii, p. 125, pl. ix, fig. 2 (1859). 

‘Twenty-one specimens, from various localities as follows :—One 9, from Poivre 
Is., Amirante Is., v-xii. 1905 (J. S. Gardiner) ; one 9, from I. des Roches, Amirante 
Is., 15. x. 1905; one 9, from Aldabra, 1907 (H. P. T.); 15 929, from Aldabra, 
1908-9 (J. C. F. Fryer); one g, two 99, from Long Island and Mahé, Seychelles 
(Cascade, about 1,000 feet), 1908-9. 

A comparison of the foregoing material with the type of the species, taken in 
Madagascar by Dr. Coquerel and now in the British Museum, places the correctness 
of the identification beyond doubt. In addition to the type of T. albipectus, the 
National Collection includes a 3 and @ from Astove I. and Aldabra respectively 
(P. R. Dupont), and a 2 from Pigaduli, Zanzibar, “on goat,” 14.x.1912 (Dr. W. M. 
Aders). 

To the specimens from Aldabra in the present series, taken by Mr. J. C. F. Fryer, 
the following interesting field-note was attached :—“ Most. numerous in December 
[1908]: attack sea-turtles, biting them between the plates on the back: apparently 
attack them in water as well as on land. Were seen flying all over the lagoon, and 
also some way out at sea. They fly strongly, and bite man also.” As already 
explained, this note must be taken as applying equally to Aegophagamyia terticeps, 
Austen (vide supra). With reference to the biting of reptiles by blood-sucking 
Diptera, it may be remembered that the typical series of Tabanus crocodilinus, 
Austen, was taken on a crocodile in Nyasaland (cf. Austen, Bull. Ent. Res., 1, 
p. 285 (1912)), while in Uganda, as is well known, Glossina palpalis feeds readily 
upon crocodiles and monitors. 


47 


ON THE REASONS FOR THE VARIATION IN THE EFFECTS OF 
FORMALDEHYDE AS A POISON FOR HOUSE-FLIES, 
By Capt. Lu. Luoyp, D.Se., R.A.M.C. (T. F.). 


The following is an account of an investigation carried out on behalf of the Hygiene 
Department of the Royal Army Medical College at the suggestion of Prof. Maxwell 
Lefroy, who, after his visit to the Mesopotamia War Area in 1916, was insistent 
on the importance of the discovery of some poison for house-flies (Musca domestica 
and allied species) that could be used indoors with safety. The work was carried 
out in the Entomological Department of the Imperial College of Science in the early 
part of 1917. Large stocks of flies which were kept breeding in two rooms were 
available for the purpose. 

Formaldehyde has been frequently recommended as a poison for flies, but those | 
who tested its effects gave varying accounts of it. Miss Lodge (1), who studied 
it in some detail, was unable to account for these variations and stated that no 
definite conclusions could be drawn from her work. An analysis of the various 
factors which might cause such variations was made, and it was found that there 
were three which play an important part. These are (1) impurities in the formal- 
dehyde, (2) the humidity of the atmosphere, and (3) the strength of the solution 
used. 

The following subjects will also be discussed, (4) the advantage of adding a bait, 
(5) the most effective mode of administration, and (6) comparison with other poisons. 

“Formaldehyde”? must be understood throughout to mean the 40 per cent. 
solution in which it is usually sold, and all strengths are given as percentages of this. 
“ Formalin,” the trade name of one brand, is often loosely used for this solution. 


Impurities in the Formaldehyde. | 

Formaldehyde is prepared by passing a mixture of heated air and the vapour 
of methyl alcohol over some heated contact agent, such as copper gauze or silver. 
The methyl! alcohol is crude or pure, but should be 90 per cent., and should not 
contain more than | per cent. of acetone. The purity of the product depends largely 
on that of the methyl alcohol, and on the freedom of the air from ammonia. 
The following substances are liable to be present: trioxymethylene, methylal, 
methyl and ethyl alcohols, acetone, hexamethylene tetramine, methylamine, and 
formic acid. Prof. Baker, of the Imperial College of Science, provided samples of 
formaldehyde to which certain of these substances had been added in proportions 
in which they are liable to occur. Solutions of these, generally in water and soaked 
up in clean sand, were exposed side by side with solutions of the control formal- 
dehyde in the fly room, and left for twenty-four hours. The numbers of dead flies 
in the immediate neighbourhood of each were then counted. Many flies left the 
table to die and no account could be taken of them. There was always a definite 
group dead in a circle of six inches radius around each jar, and the comparisons 
were based on these. In order to obviate the influence of the varying numbers 
of flies on different days, all numbers are given reduced to percentages of the controls. 
All decisions are based on a number of tests, with the positions of the various solu- 
tions and the controls interchanged. 


48 CAPT, LL. LLOYD. 


Of the impurities tested, hexamethylene tetramine, methylamine and formic 
acid were found to be of importance, and these only will be discussed. The effects 
of the presence of the other substances mentioned above were investigated and found 
to be neutral, or of very little importance. 

Hexamethylene tetramine, (CH,),N,. 

This substance is produced by the action of ammonia on formaldehyde. It is 
liable to occur in the solution owing to the presence of ammonia in the air during 
manufacture. It breaks down with the formation of methylamine, CH, NH,. 
Formaldehyde containing these bodies has a yellowish colour and a fishy odour, 
both these characteristics being intensified by the addition of a strong alkali. These 
substances are distasteful to flies. A sample containing 5 grms. of hexamethylene 
tetramine added per litre was tested against a control. The residue from the manu- 
facture of this substance, containing ammonia and methylamine, and some methyla- 
mine hydrochloride, with and without an alkali to set free the methylamine, were 
also tested with the following results. 


TaBLeE I. 
Showing the Influence of Hexamethylene tetramine in Formaldehyde. 


Soluti | Percentages of flies | Average | Total flies 
olution | killed | of | killed 
Formaldehyde control, 10% 100 in each case 100 324 
Formaldehyde with 5 grms. 
hexamethylene tetramine 
added per litre, 10% T45/430; 932071. obi 85 267 


Tas_e II. 
Showing the Influence of Methylamine in Formaldehyde. 


Boise Percentages of flies killed (actual 
numbers in brackets) 
| 
Formaldehyde control, 109; 100 (50) 100 (20) 
es with 2°5% hexamethylene 
tetramine residue 10% 36 (18) 
#} with 1% hexamethylene | 
tetramine residue 10% 65 (13) 
os with 1% methylamine 
| hydrochloride 10% | 40 (8) 
re with 5% methylamine hydro- 
chloride and 0:2% KOH 
10% 30 (15) 


3 with 1% methylamine hydro- 
chloride and 019% KOH 
i ae ae ae a pou 30 (6) 


A sample, naturally containing these substances in unknown proportion, with 
a pale yellow colour and fishy odour (previously unsealed bottle from the store 
of the Imp. Coll. Sci.) was tested against a colourless control with a clean odour. 


FORMALDEHYDE AS A POISON FOR HOUSE-FLIES., 49 


The results of the tests are summarised in the next table. In nearly every case 
the purer sample proved the more effective. No suggestion can be given for the 
purification of formaldehyde of this nature, and it should be avoided if any other 
is obtainable. 


TABLE IIT. 
Comparing a Yellow Formaldehyde with Fishy Odour (Methylamine) with a Clean 
Sample. 
Soluti Percentages of flies Average Total flies 
kcal | killed % killed 
Formaldehydecontrol, 10% | 100in each case 100 | 1509 
a impure, ae | ted. Gi, 96, -89, 230, 36; 
| 17, 68, 58. 89 1118 


| 


Formic acid, H’CO’'OH. 

Formaldehyde is prepared by the dehydrogenation of methyl alcohol: CH,OH + O 
= CH,O + H,0. Theoretically in the process there is no formic acid formed. The 
samples of formaldehyde examined, however, were all found to be acid. Three 
which were titrated were found to be +24, +39, and +60 acid (Eyre) respectively. 
(A fluid ““ +a acid ” requires x ccs. of normal alkali to make one litre neutral.) 
The acid present was proved to be formic by a test suggested by Dr. A. M. Whiteley. 
A few ccs. of the formaldehyde are shaken up with lead carbonate and quickly 
filtered. If formic acid is present crystals of the sparingly soluble lead formate 
separate out in the filtrate after a short interval. The crystals are minute, very 
characteristic needles. On the assumption that formic was the only acid present, 
these samples contained 1°1, 1°8, and 2°76 grms. of formic acid per litre respec- 
tively. When the dilute poison 5-10 per cent. formaldehyde is made up the acid 
would be present in amounts varying from ‘0055 to ‘0276 per cent. Hven these 
small quantities are detected and disliked by flies. 


TaBLe IV. 
Showing the Deterrent Effect of Formic Acid in Water. 


euriian No. of Av. flies | Av. flies Av. percentage 
tests | tasting | drinking which drank 
Tap water control 2 35 24 69 
Formic acid ya'8 1 33 | 3 10 
i 05% 3 32 | 15 47 
ag 02 :049 4 36 20 54 
in 01 005% 3 36 | 18 50 


Temperature 77°9° F. ; relative humidity 57%. 


Testing saucers, two inches in diameter, were filled with solutions of formic acid 
in water and covered with discs of blotting paper to soak up the fluid. These 
were then exposed one at a time in the fly room and watched for five minutes, a count 


50 CAPT. LL. LLOYD. 


being taken of the numbers of flies which visited and tasted the solution and then 
left, and of those which remained to drink. Controls were made with tap water. 
A fly only “ tastes ’’ when it presses its proboscis one or more times against the wet 
surface, and quickly withdraws it. It “ drinks ’’ when the proboscis remains pressed 
against the surface for a few seconds or longer. Apart from the count, an impression 
may also be gathered from the behaviour of the flies. On a fluid which they find 
attractive they cluster, but on an unattractive one there are never more than two 
or three flies at one time. The total number of visiting flies was about constant 
in each test. 

The disadvantages of the presence of formic acid in formaldehyde are shown 
by the following tests, where a solution containing 12 grms. per litre was tested 
against the one containing 1°8 grms. per litre. In ten tests the less acid solution 
killed twice as many flies as the more acid one. 


TABLE V. 
Showing the Disadvantages of the presence of Formic Acid in Formaldehyde. 


Soluti Percentages of Av. per cent. Total 
ete toon flies killed killed killed 
Formaldehyde 1:8 grms. | | ) 
ECF OE | 
per lit. 10% | 100in each case 100 808 
39 12-0 grms. | | | 
He OOF E. | | 
per lit.10% | 54, 78, 54, 27, 34, 38, 34, | | 
51, 43, 50 | 50°6 | 366 
j 


Flies are less readily repelled by dilute alkaline than by acid solutions, and will 
drink solutions containing 0°1 to 0°2 per cent. caustic potash, or 0°07 per cent. 
lime, as readily as water. There is possibly even a little attraction in it. The fol- 
lowing Table VI shows this, the data being obtained in the same way as those in 


Table LV. 


Tasie VI. 
Showing that certain weak Alkaline Solutions are not deterrent. 
| | 
No. of Av. flies {| Av. flies Av. percentage 
Solution tests tasting drinking which drank 

| 
Tap-water control 5 38 ol 83 
KOH2% 1 35 0 | 0 
» 0°3-0°4.% 2 31 14 | 45 
»  0°1-0:2% 2 | 34 30 | 91 

Ca(OH):, 0°14% (lime | 
water: 2 37 20 54 
» _ 0°078% 2 | 33 20 61 
» 007% 3 | 37 30 81 


Temperature 


77° F.; relative humidity 57%. 


FORMALDEHYDE AS A POISON FOR HOUSE-FLIES. 51 


It is therefore advisable always to add as much alkali to the formaldehyde as 
possible, without deterring the flies, to neutralise any formic acid which may be 
present, or any which may be formed by oxidation during the exposure of the solu- 
tion. Table VII shows the benefit of the procedure. 


TaB_eE VII. 
Showing the Advantages of adding certain Alkalis to Acid Formaldehyde. 


ye | Percentages killed (actual 
Solution, 10% in each case numbers in brackets) 


| 


Control formaldehyde (12°5 grms. H-CO-OH per lit.) 100 in each case 
+50% lime water in dilution 283 (155), 241 (82) 


» +K,CO,, 15 grms. per. lit. 300 (36), 268 (147), 270 (92) 
(equiv.). 
+(NH,),CO;,10 grms. per lit. | 
(equiv.). | 68 (37), 82 (28) 
Control formaldehyde (1:8 grms. H-CO°OH per lit.) 100 in each case 
. +100% lime water indilution | 76(47), 90 (923) 
* +P. . ss * - " | 114 (269), 104 (312), 142 (57) 
” apa POG CO. | 125 (50) 
> + 1% NH,0OH | 50 (20) 
Control formaldehyde (1:1 grms. H:CO-:OH_ per lit.) 100 in each case 
= +502, lime water in dilution | 131 (297), 124 (31) 
9 +25% ” 2? oh 2” 110 (28) 


Ces cepa ye cena 


It will be seen that the addition of the alkali improved the effect in every case 
except when the whole fluid of dilution was lime water, which made it too strongly 
alkaline, and when ammonia was used, methylamine being formed in these cases. 
Time water is recommended in practice in preference to other alkalis because it is 
a weak solution of definite strength. It is prepared by allowing cold water to stand 
over lime, slaked or unslaked, for a few hours and then filtering the solution before 
use. Such a solution contains 0°14 per cent. Ca(OH),. If made with hot water 
the solution is weaker. The maximum amount, half the total fluid of dilution, 
should always be used to counteract the formic acid which would otherwise be 
formed after dilution, during exposure to the air, or by the bodies of dead flies 
falling in. In a trap which will be presently described, but an imperfect model, 
which was exposed in the fly room, this was very clearly shown. The fluid in the 
trap was 7°5 per cent. formaldehyde, 35 per cent. lime water (equivalent to the acid 
present), and 5 per cent. sugar. On the first three days this trap killed 1,800 flies, 
and then became ineffective. Four flies had crept into the poison, and when this 
was titrated it was found to have become +2 acid. An equivalent of KOH was 
added and it became again effective. 


Humidity of the Atmosphere. 
A test was made at a temperature of 96° F., and the poison was found to be 
effective. There is no reason apparent why it should not be effective in a tropical 
country. The humidity of the air, however, plays a considerable part, as indeed 


5? CAPT. LL. LLOYD. 


it must with any poison which is administered as a drink. In wet weather flies 
have less inclination to drink and there is more moisture available, so that they 
are less likely to discover the trap. Experiments were made to test this. 

(i). A jar with an area of 2°5 sq. in. was filled with sand wetted with a 10 per cent. 
solution of formaldehyde without any addition. It was placed in the centre of a 
dish of dry sand, area 95 sq. in., and this was placed in a greenhouse containing a 
large number of flies. No water was available and the atmosphere was dry. After 
twenty-four hours 1,560 dead flies were counted on the surface of the sand, or within 
a few inches of the dish. The experiment was repeated with approximately the 
same number of flies available and the same atmospheric conditions, but in this 
case the sand in the surrounding dish was wetted with water, equally with the 
poisoned surface. After twenty-six hours only 20 flies were found dead, the majority 
having obtained their water from the unpoisoned surface. 

(1). This series was carried out in a glass cylinder 24 in. high and 10 in. in diameter. 
Wet and dry bulb thermometers were included. The formaldehyde (10 per cent. 
in water) was supplied soaked up in sand, area 2°5 sq. in. The duration of each 
experiment was twenty-two hours. The air was in no case quite dry, as the amount 
of moisture given off by the poisoned surface is considerable in this confined space. 
It was found that the formaldehyde was effective with a relative humidity of 76 
per cent. and 72 per cent., that its effect was about halved with a relative humidity 
of 81 per cent., and that above 90 per cent. it was much reduced, available water 
being present throughout in this case. The results are given in Table VIII. From 
these it may be concluded that liquid poisons will not be very effective if 
much other available moisture, such as dew, is present. Formaldehyde works 
well in a room where there is a wet sink, as was the case in the fly room where these 
experiments were performed. It would be useless, however, to place the poison 
in the sink or immediately over it, though it will be found very effective a few feet 
away. Flies are usually going about looking for food, and seem to taste all moisture 
they come across in their wanderings, either in the hope of its containing food or 
simply for the sake of the drink. Their capacity for utilising solid food depends. 
on whether the crop contains fluid with which to dissolve it. 


Tase VIII. 
Showing the decreasing effect of Formaldehyde with increasing Humidity. 


| 
Conditions in Cylinders Temp. | Rel. Humid | Total flies | Perooniage 
| | | 
CaCl, added to dry the air . | Tor" | 7 pay 229 94 
Air not treated | TaOAT we 76% | 125 89 
Wet blotting paper as base | da°0° 81% 66 47 
Wet blotting paper as base and top | 76°3° | 93%, 90 17 


(ui). Experiments with baited formaldehyde under saturated conditions were 
made. It was found that the baits about doubled its efficacy, but the results were 
still very unsatisfactory. The details are given below in Table IX. Sugar in water 
was the bait and the formaldehyde was employed in various strengths, being supplied 


FORMALDEHYDE AS A POISON FOR HOUSE-FLIES. 53 


in two-inch saucers with blotting-paper covers. Each series was continued for 
two days and the cyliriders were kept as wet as possible, though in several cases 
they dried somewhat towards the end, with a corresponding increase in the death- 
rate. The controls were on a dry table with muslin tops to the cylinders, the 
relative humidity being about 76 per cent. The fourth experiment in Table VII 
is taken as a control for the saturated conditions with unbaited formaldehyde. 
To obtain a proper estimate the percentages dead at the end of twenty-two hours 
should be studied. Another series carried out in precisely the same way, but with 
bread-water as a bait instead of sugar, gave very closely similar results and will 
therefore not be detailed. 


TABLE IX. 
The effect of Barted Formaldehyde compared under Wet and Dry Conditions. 
Bed Se ae 5 at SER Se Oe es ac 
| Percentages dead after : 
Solution _ Conditions | No. of flies _K-—_, —_____—_—____- 
5 hrs. 22 hrs. | 48 hrs. 
No bait, formaldehyde 10%, | wet 90 | 17 
Sugar 5% As ne dry 38 | 32 97 100 
* ” » » wet 34 | 3 35 79 
a d TH, | dry Ce ee 9] 100 
e. Be a ~ wet 28 3 Zo 69 
ar ¢,: . 50% | dry 21 43 100 100 
Nes A .. wet Ce as 4 22 78 
* 4 . es ey a diy 19 | 2) 79 | 100 
a “A te i oe wet 17 | 0 6 | 82 


Humidity and the use of formaldehyde on ships. 

Lt.-Col. W. Byam, R.A.M.C., reported that while he found formaldehyde very 
effective on board ship, it failed to be of use in Egypt. This is in accordance with 
the variations in the accounts of its utility. That used in Egypt was possibly too 
acid. Experiments were devised to discover what its effect would be on board ship 
where the atmosphere is moist, but not necessarily saturated, and where most of 
the available water is salt, owing to all the woodwork being saturated with this 
substance. 

(1). A cylinder was washed inside with 10 per cent. salt solution, flies were intro- 
duced and blotting-paper soaked in the salt solution placed above and below; a 
shelter was erected over the poison dish to prevent the salt water dripping in. A 
control was set up, fresh water being used instead of salt solution. It was found 
that the flies were poisoned about twice as rapidly in the one with salt water as in 
the one with fresh water. 

(2). This was carried out in cylinders, a bait being used in the poison which was 
soaked up in jars of sand (area 2°5 sq. in.) that were placed in dishes of sand (area 13°5 
Sq. in.) wetted in one case with 10 per cent. salt solution and in the other with water. 
The atmosphere would be very moist from the vapour given off by the wet sand, 


54 CAPT: LL. LLOYD. 


TaBLE X. 
Showing the Effect of Formaldehyde when available Water is strongly saline. 


Percentages dead. after : 


Solution Conditions Total flies - : a 
| j 
| 
3 hrs. 28 hrs. | 48 hrs. 
| 
Formaldehyde Saturated with 
10% in water salt solution | 29 14 69 93 
Formaldehyde | Saturated with | | 
10% in water | fresh water | 27 4 | 39 78 
| 


and the conditions would approach those on board ship when it is not actually 
raining. The results, which are given in Table XI, are very striking, as the flies 
died rapidly when the available water was saline, and slowly when it was fresh. By 
further experiment it was found that the flies drink as readily as water a solution 
of 1 per cent. salt, that stronger solutions deter them, while 5 per cent. and upwards 
repels them entirely. Formaldehyde properly neutralised and baited will be found 
very effective on board ship. 


TABLE XI. 
Showing the Probable Effect of Formaldehyde on Board Ship. 


| 


| Percentages dead after : 
} 
| 
| 


Solution Conditions Total flies = oe 
te Ries fl Sekai, 24 hrs. 
| | 7 
| 
eae Ae | <i PEE Oe ae | 
Relies te available | 94 epee: 43 96 
bos ater 5 3/2” | Fresh water 
sugar 5% / available 38 | 3 | 2) 63 
| 
| | | | 


The Strength of Formaldehyde to be used. 


The question as to the most effective strength of formaldehyde was approached 
trom two points of view; firstly, the stronger the solution the less likely are the 
flies to drink much ; secondly, a little of a strong solution, or a large quantity of a 
weak solution, will kill them. ‘These will be discussed under the headings of “‘ The 
Deterrence of Formaldehyde” and “ The Toxicity of Formaldehyde.” The results 
were then combined, giving the optimum concentration. At the same time control 
experiments were made in cylinders as a check to the conclusion drawn. 


i | 
a 


FORMALDEHYDE AS A POISON FOR HOUSE-FLIES. 


The Deterrence of Formaldehyde. 

The deterrence of formaldehyde was determined by the same method as that 
employed with formic acid, already described. Each series of different strengths 
was exposed, one at a time and in quick succession, in the same spot where the 
numbers of available flies were about constant. The numbers which only tasted 
and the numbers which drank were counted and the percentage of the latter to the 


90 


Oh Lo" 
etl yf retreat ain 
a sa “oe: eae he 

Yl l nereobel d 
yp 
SE 


10 95 90 85 6:0 75 7:0 65 6055 §0 45 40 35 3:0 25 20 15 10 
Percentage of strength in dilution. 


@ 
2) 


~ 
° 


a 
°o 


oO 
° 


> 
ro) 


1) 
oO 


Percentage of tasting flies which drink, 


1a) 
°o 


io 


F  secoscccceveceee Lormaldehyde and water, and control of water. 
aS ¥ 5 59 % lime water £ ,, 00 % lime water. 
oe ee ee ” ” 5 5% sugar 3 5 of sugar. 


50 % lime water and 5 % sugar. 


Diagram 1. Showing the deterrence of acid formaldehyde and its remedy. 


former gives the factor to the solution. Controls were made with water alone, 
with 50 per cent. lime water, and 5 per cent. sugar solution, and each series was 
checked by one or more repetitions. The time allowed to each exposure was five 
minutes. The following solutions were thus studied: formaldehyde in water ; 
formaldehyde in 50 per cent. lime water ; formaldehyde in sugar solution ; formal- 
dehyde in 50 per cent. lime water with sugar. The detailed results are given in 


56 CAPT MLL: LLOYD: 


Table XII, and in Diagram 1. In the diagram the points represent the percentages 
of the tasting flies which remained to drink. The bottom dotted line represents 
formaldehyde and water, and shows that any mixture of these is deterrent as com- 
pared to the water control, of which an average of 82 per cent. of the tasting flies 
drank. The 10 per cent. formaldehyde deterred from drinking nearly 82 per cent. 
(67 out of 82) of the tasting flies which would have drunk had there been no formalde- 


Taste XII. 
Showing the Deterrence of Formaldehyde, and us Remedy. 


| | | | | 
No.oi/Total| No. of flies | No. of flies | Percent. of 
tests flies tasting drinking drinking to 
; | tasting flies 
Solution | 
| | | 1). | 
| Av Max Mir Av Max) Min Av |Max/ Min 
Water controls | 8 |. 263 | 33 ae P26 eer 1185 | 20"! Sasa ree 
,, formaldehyde 10%! 6) | 227 |S SO COR ras Peles ita! Sel ee eal ees 
* i‘ 75%, 6 | 190 | 32 | 35) 27 | 7) 13) 0} 16) 38} v 
i f 50%!) | 6 195 | 32° 1°38 126.) | es oan Saag 
; 7 25% 6 -|} 297°| S50 49ekoG, | 92/97 Nia BOr aSrlise 
. ds 20% | 4. | 148 ]-39-|-634) -24en ol hoot sel Ou AT 9) Be 
¥ : 15%, | | 4} 151 | 38 | 47 | 29) |° 26.) 34 1G: | ee ee 
; ‘ 10% | 2 715.1 37 | 38. 37) | 230318 26) | ised ee 
33 - me} ule 74. | 37 | 28 | 36.) 30 | Tals 7 30) 2 san em 
Lime water controls 50S, 3 112 | 37 | 39 | 35 | 30 | 31 | 29 | 80 | 86 | 72 
,. formaldehyde 10% 4 }| 186 | 34. | 42°°255" Sed 1 Gu 29) eu cae 
a * 75% a0} 121-30" |. 407723) tes os) ae 
ie ¥ 5-09, 4 |} 122 | 30| 33 | 28)) 16 | 2E Uh oad ee ee 
- . 2:59!) | 14 [| 114) 28 | 320s) 265) 20 23 a aaa eel ae 
‘ - B0% bo D264} B24 -BB- 1B BB125 4) abel ee 6) Be 
‘ P 15% 2.4 HO /°35 | 89-1 BLS) 28:1 27 7 20) G6 ae Sao 
Water, sugar controls wy ae wl | 45 | 45 | | 40 | 89 | 
, formaldehyde 10% Bh 6 | BS |. 45°30 Tae) Ol ls axl at 6 cae 
ss 5 T5O,, 2 63 | 31 | 36 | 27 | 17 | 23 | 11 | 52 | 63 | 41 
: ? Oo, 2 7. | 35 | 38 |-33" 22 | 284 2h | 62°|.69:) ae 
> ? Deb, 2 66 9/933"| 35°| 31-28. 31; | 265/86 | 88 | BA 
Lime water 50% | | | 
sugar 2 a | | | 
, formaldehyde 10% | 2 60) 30 | 804 30°] Ia tre 47 eae 
99 : 1 ae a 72.| 36 | 41 | 31 | S429 20.) 68) Fes 
: “ BUcG, | 2 60 | 30 | 35'| 25 | 23 | 27 | 20 | 78 | 80 | 77 
, es oy Se 52.126 | 27 | 25 | 22) 24) 21 eG eae 
| | | | 


Temp. av. 77° F. (78:9-76:0°). Rel. humid. av. 50% (58-39%). 


hyde present, and so on. The line of dashes represents the percentages which 
drank the formaldehyde in 50 per cent. lime water. It is higher than the previous 
one because, while the deterrent influence of the formaldehyde remains as before, 
that of the formic acid is removed. It also is always below the water and lime water 
controls. The next line of dots and dashes represents formaldehyde and 5 per cent. 
sugar. without lime water. The presence of the bait has partly counteracted the 


FORMALDEHYDE AS A POISON FOR HOUSE-FLIES., 57 


deterrence, so that at 3 per cent. it is equal in its attraction to the water control, 
and at 2°5 per cent. approaches very near to the control of sugar and water. The 
continuous line represents the mixture which is finally recommended, 50 per cent. 
lime water and 2°5 per cent sugar. It is equal to the water control at 5°5 per cent., 
and then becomes more attractive than water because the flies recognise in it a food, 
while the formaldehyde is not repellent enough in weak solution to deter them in 
large numbers. At 5 per cent. only 12 per cent. fewer flies drank than would have 
done so if the solution were sugar and water only. At 2°5 per cent. formaldehyde, 
only 3°5 per cent. declined to drink. The neutralisation of the formic acid and the 
addition of the bait have at this point nearly counteracted the deterrent action of 
the formaldehyde. It is of considerable interest to notice that the lines which 
represent the formaldehyde with and without lime water are parallel in pairs, while 
those with and without sugar converge in pairs at 2°5-1°5 per cent. This shows 
the influence of the presence of formic acid in very dilute solutions. The formalde- 
hyde used proved on titration to be + 60 acid (Eyre) which is equivalent to 2°76 
grams of formic acid per litre of formaldehyde. At 5°0 per cent. dilution this is 
0°014 per cent. formic acid, and at 2°5 per cent. it is 0°007 per cent. formic acid. The 
presence of sugar is sufficient to counteract the latter but not the former. A factor 
which has not been taken into consideration is that the formaldehyde vapour may 
be deterrent and that fewer flies may taste stronger than weaker solutions. This 
was not apparent in the open, but showed itself when the experiment was repeated 
in enclosed spaces. There was then progressive deterrence as the strength of the 
solution increased, as was shown by the smaller percentage of the available flies 
which settled on it. The curve of this deterrence is represented by the dotted line 
in Diagram II. Consideration of it will show that its fall is so gradual that it would 
not affect the argument even if its influence were as powerful in the open as in the 
enclosed space, since the influence of the other factors is so much more potent. 


The Toxicity of Formaldehyde. 

In determining this factor, the solution used was 50 per cent. lime water and 2°5 per 
cent. sugar, only the percentage of formaldehyde varying. About 50 flies were 
enclosed in each of several glass basins of capacity of 600 cc. The basins were then 
placed over saucers containing the poison and a count was taken of the flies that 
tasted and those that drank. In the first series the time allowed was fifteen minutes, 
but it was found that all the flies that intended to drink did so in the first five minutes, 
and the exposure was subsequently limited to this time. The poison was then re- 
moved and the edge of the basin slightly raised to allow air to circulate. At the end 
of one hour and of two hours the numbers lying inert were counted, the difference 
between the two counts representing the recoveries, as formaldehyde seems to act 
as an anaesthetic, and its effect in weak solution, or when a very little is taken, is 
sometimes only temporary. No recoveries were noted after two hours. All the 
flies were then anaesthetised and counted. Four series of each percentage were thus 
made. The complete results are summarised in Table XIII. This shows the total 
and average numbers of flies exposed to each percentage of formaldehyde used (from 
2°5 to 20 per cent.) ; the average, maximum and minimum numbers of flies which 
tasted ; the average, maximum and minimum percentages of those which drank 

(681) r 


58 CAPT, LL, LLOYD. 


compared to those which tasted ; and the percentages of those temporarily and 
permanently incapacitated compared with those which drank. The calculation of the 
numbers of those which died against those which drank is more satisfactory than 
calculating them against those which only tasted. The numbers thus obtained are 
frequently above 100, as at a strength of 5 per cent. and upwards the amount taken 
by a fly in its taste is often enough to kill it. Reference to the table will show that 
not only the averages, but also the maxima and minima, run in series corresponding 
to the strength of the poison, allowance being made for occasional variation due to: 
experimental error. The percentage of those which died compared to those which 
drank gives the factor of toxicity to each strength of formaldehyde. These factors 
are plotted in Diagram 2 and are represented by the line of dashes. The toxicity 
increases rapidly from 15 per cent. upwards, and falls rapidly from 5 per cent. down- 
wards, the rise from 5 per cent. to 15 per cent. being gradual. The dot and dash curve 
is taken from Diagram 1, with the addition of the factors for 15 per cent. and 20 per 
cent. strengths which are taken from Table XIII. This curve represents the per- 
centage of tasting flies which remain to drink a solution of formaldehyde containing 
50 per cent. lime water and 2°5 per cent. sugar. It is the curve of deterrence. By 
combining these two curves a hypothetical curve is constructed which is represented 
in the diagram by the continuous line. For example with a strength of 10 per cent. 
formaldehyde exposed in the open, 47 per cent. of the flies that taste it will remain to: 
drink. Some however will absorb enough poison in the taste to kill them, and the 
number which finally die is not 47 per cent. of the tasting flies but 133 per cent. of 47, 
that is, 62°5 per cent. of the flies that taste die. The relative effects of different. 
strengths of formaldehyde, as represented by the curve, were thus found. Itis a natural 
curve with its maximum at 5°0-6°0 per cent. formaldehyde, and this is therefore the 
optimum strength to use in practise. The straight dotted line in the same diagram 
represents, as explained above, the deterrence of the vapour at different strengths in 
enclosed spaces. Its tendency is to recommend the use of as weak a solution as 
possible, but the decline in the effect below 5 per cent. is so rapid that this factor could 
play no part in the decision as to which strength to use. 


TaBLe XIII. 


| | | | | % to drinking flies 
At ; | | | % of flies | % of tasting 
Solution =| ln ,.,| Av. | tasting | flies which |— a. 
50% lime \Tests |Total) jo. | s eee de 
ee ae | no. | flies | Aies | | Downin| Dead after 
| ie fea gyi ee ee ee rn Tee 2 hous _ 
|Av. Max. Min.|Av. Max. Min. Av. Max. Min. 
Formald. 20% | 4 | 154] 39] 483 67 30/20 25 15 | 229 | 219 300 125 
o 165% | 3 (188) 44} 41 53. 85 \| 80.3625.) 146 9) 1437 150 "140 
2 1%, 4 | 220; 55 |} 53 55 45 | 54 67 48 136 133156. 110 
a 75%! | 4 | 1821 45159 69 49|68 96 63) 111 | 112 136°. 87 
2 50% 4 | 219! 55) 61 65 53 | 86 93 79 106) 10G°"°12T ' 86 
. 275% 4 | 174, 43 | 66 80 54/9196 87): 61.4 39 962) 23 


FORMALDEHYDE AS A POISON FOR HOUSE-FLIES, 59 


cS 
_~ 
_—— 
=, 
= 


Percentages of Fiies 


12:5 10 7°5 
Percentage strength in AGilution. 


—..—..—.. Flies tasting fluid which remain to drink. 

Percentages killed of those drinking. 

Relative effect of various strengths. 

rae eae eee Deterrent effect of formaldehyde vapour in enclosed spaces. 


Diagram 2. Showing the most effective strength of formaldehyde. 


The Optimum Concentration. 


The conclusion drawn from the experiments just described was checked by the 
following series of further experiments. In each series concentrations of 10 per cent., 
75 per cent., 5°0 per cent., and 2°5 per cent. formaldehyde, with 50 per cent. lime water 
and 2°5 per cent. sugar, were placed in saucers with blotting paper covers and these 
were exposed to flies in cylinders (10 in. diam., 8 in. high). Each experiment was 
continued for twenty-four hours, and counts were made at intervals of the flies lying 
inert. Seven series were completed, and about 100 flies were used in each cylinder. 
The summary of the whole is given in Table XIV. In this are shown the average 
number of flies in each cylinder, and the average percentages inert at the end of 
various intervals. These results are plotted in Diagram 3; the dot-and-dash line 
represents 2°5 per cent. formaldehyde; the continuous one 5 per cent.; the line of 
dashes 7°5 per cent.; the dotted line 10 per cent. Although recoveries are only 

(681) 2 


60 CAPT, LL. LLOYD. 


indicated in the case of 2°5 per cent., they occurred occasionally also in the case of 5 
per cent. and 75 per cent. strengths. This method confirms the conclusion that the 
most effective strength to use is 5 per cent., then 7°5 per cent., then 10 per cent., while 
2°5 per cent. 1s the least effective. The factors of effect of these concentrations had 
been found to be 80°3, 7671, 62°5, and 33°5 respectively. 

TABLE XIV. 


Showing the Death-rate of Flies exposed to various strengths of Formaldehyde. 


Formaldehyde No. of Av. no. . ; 
strength Vouie flies Av. percentages inert after : 
EO: Ba : 
1 hour | 3 hours | 4 hours | 5 hours | 24 hrs. 

a 81 arr ae wee: 92 
1096 7 92 4a | 53), |. 63 |. 6p 93 
5-0% 7 98 St ee oer oe 96 
25%, di 96 42D hd ASE Ca 53 89 


| 
g pests 
— 
oe ee Lee | 
a 
=, = 
a 
a 
cS e 
oe ae e - 
oe .° 
st 
x 
+ 
e— 
i - 
a | er ae 
_ ry 
=-_ el? e ee 
= e? . 
c a° 
ae ° } 
4 °° e 
: 2° 
hs °°? 
. 
f . 
or 


60 


~ 
© 


ag Fae p 
$s 

60 

y 

v 

S 

5p 

n 

S 

c 40 
v : 
he i 
Y 
Q 30-4 


Le) p op 7G "gh em ame Mete ia Tht) ie 


! IS) 14S 1G) 7) 16 MISE 2021 Wee (26. es 
/nterva/ls of time in hours. 
One 2S Formaldehyde 10 % 
ie eee 3 13% 
9 5°0 wf 


0 
"Fh O 
— +? au 6 6 «<= © 6 9 2°5 Yo 


20 


sO 


Diagram 3. Showing the relative strengths of formaldehyde. 


FORMALDEHYDE AS A POISON FOR HOUSE-FLUIES., 61 


Advantages of adding a Bait. 

Reference to Table XII and to Diagram 1 will show that to get the best effect from 
the formaldehyde it is necessary to add some bait to counteract the deterrent effect. 
By this means a larger number of flies are induced to drink. Sugar is the most 
attractive and most permanent bait, and 2°5 per cent., or half an ounce to the pint, in 
the dilution is sufficient ; to make it stronger does not make it more effective, while at 
1 per cent. it begins to lose its influence. An advantage of this substance is that it 
has no interaction with the formaldehyde. Glucose, maltose and lactose have also 
been tried, but are less effective than cane sugar. Molasses will be found effective. 
An infusion of hay, boiled and strained, and a cold infusion of horse-dung have been 
tested and found to be effective, but less so than sugar, and they are not recommended 
for this reason. 

The formula to be employed is :— 
40 °, Formaldehyde, 50-60 ce. ag i 1 fluid ounce. 
Filtered lime water, 500 cc. .. ae i! 4 pint. 
Sugar, 25 grams Ns my a ounce (dessertspoonful). 
Water, add to make 1,000 ce. . * add to make 1 pint. 


Method of Administration. 

It is necessary to protect the formaldehyde from the action of air, and to prevent 
flies from falling into it, both of which factors make it acid, and soon cause the lime 
water to be neutralised. A simple form of trap has been devised for this. The 
poison is placed in a bottle, and the mouth is closed by means of a platform of absorp- 
tive material from the centre of which a stem of the same material passes down into 
the fluid. The top is wetted with the fluid at the commencement and is kept wet by 


rol— bo 


Fig. 1. Form of trap for 
use with formaldehyde. 


capillary attraction. Such an arrangement may easily be made from blotting paper. 
A circle of the paper two inches in diameter is cut out, and a small slit is made in the 
centre. ‘Two strips of the paper of the same width as the slit are then passed through 
it and the ends are doubled outwards and downwards. The strips are then drawn 
down till the doubled-back portions rest on the platform, and the trap is ready for 
use (see fig. 1). 


62 CAPT. LL. LLOYD. 


This form of trap may be used under cover or outside when it is not raining. The 
paper should be renewed every third or fourth day in any case. A more permanent 
form may be made from plaster of Paris or some other durable material, porosity 
being an essential. The stem in this case is circular and should be in one piece with 
the top, or fit into it very tightly, otherwise the fluid will not rise. It may be 
necessary to soak this trap im water occasionally and dry it. 

These traps have been submitted to the very severe tests of being placed near 
dishes of the food which has been found most attractive to house-flies, casein, banana 
and bread, mixed and wetted. One trap with the top made of plaster of Paris was 
placed about a foot away from a large dish of this material, and in three days poisoned 
1800 flies, which were found on, or within a few inches of it. After further neutra- 
lisation it became effective again and remained so for ten days, about half the original 
fluid (200 cc.) having been used. Another one, with the top made of filter paper two 
inches in diameter, was placed in a corner of the fly room, and was active for ten days 
until it was removed, the paper top having been renewed once; 1,560 flies were found 
dead around it. A third one with a paper trap was still active at the end of a month. 
Whether a trap is still effective may be determined by sweeping away the dead flies 
from its vicinity. If no more flies accumulate, and there are still flies about, the 
fluid has probably become acid and should be renewed, as it will not be found worth 
while in practice to neutralise it again. The traps should be placed wherever flies 
are numerous, especially near rubbish bins, in stables, near manure heaps, in kitchens 
and latrines. Observation will show which are the best positions. 


Comparison with other Poisons. 


Formaldehyde as a fly poison was not tested against sodium arsenite, but it has 
two great advantages over it : it is safe and it is more economical. With 2 per cent. 
sodium arsenite solution 10 per cent. sugar is recommended, while formaldehyde only 
requires a quarter of that amount. The use of sodium arsenite would always be 
dangerous to man and to domestic animals, especially if issued to the general 
public. That the poison used shall be safe in any hands is a great desideratum. 

The action of formaldehyde was tested against that of sodium fluoride, recently 
recommended as an effective fly poison (Jackson and Lefroy). Formaldehyde (5 
per cent.) and sodium fluoride (1 per cent.) in saucers were placed in two separated 
halves of a deep glass dish, equally lighted and heated. No baits were used in either. 
After twenty-four hours in the formaldehyde section 650 flies were dead as against 234 
dead in the sodium fluoride section. As more might have escaped from the section 
containing the fluoride than from the other section, a control comparison was made 
with cylinders of flies, four containing as poison | per cent. sodium fluoride with 3 per 
cent. sugar, and four containing the formaldehyde mixture recommended above. 
Notes were made of the death-rate and it was found that there was no appreciable 
difference in the actions of the two poisons. The results are summarised in Table 
XV. However from the first experiment it is seen that flies poisoned travel less from 
the formaldehyde than from the fluoride, which is an advantage. Authorities differ 
as to whether sodium fluoride is a poison to man, but as it is used in food preserving 
it is certainly not a very dangerous substance. 


FORMALDEHYDE AS A POISON FOR HOUSE-FLIES, 63 


TABLE XV. 


| 


Aver. percentages dead after : 
No. of Aver. ad 
Solution. tests. | flies. | | 
1 hour 4 hours 24 hours 
} 
_ a A : hes 2 
1% sodium? fluoride, 3% 
sugar ieee ae 57 76 94 
5% formaldehyde, 50% | : 
lime water, 3% sugar 4 | 74 6G: {| 74 96 
Summary. 


The effective action of formaldehyde as a fly poison depends on the freedom ol 
the-exposed fluid from formic acid, and, to a less extent, from methylamine. That 
used for fly poisoning should therefore be colourless and free from a fishy odour, 
and a weak alkali, in slight excess, should be added in dilution to neutralise any 
acid present and that which will be formed during exposure. The formula recom- 
mended is :— 


40% Formaldehyde .. + aa s be em oily 
Clear lime water oy oe 1 rm al - 50% 
Sugar .. ne Zs 2% ae “ee y Pe OF 
Water .. ; re: add to make 100 


This should be exposed in a trap which will protect it from the air and will prevent 
flies from falling in. Formaldehyde, like any other stomach poison for flies, will 
be most effective under dry conditions. 


References. 
1. O. C. Lodge, Proc. Zool. Soc. Lond., Sept. 1916. 
2. Thorpe, Dictionary of Chemistry, on formaldehyde. 
3. Mally, C.W. “On the use of poisoned bait for controlling the house-fly ; ” 
S. African Journal of Science, June 1915. 


4. Jackson and Lefroy, “Some Fly Poisons for Outdoor and Hospital Use ;”’ 
Bull. Ent. Res. vii, p. 327. 


Lees, 7 aa ; 
‘if Lee 
» 4? 


AFRICAN APHIDIDAE.—Parr IV. 


By Frep. VY. THEOBALD. 


The following new species of Aphids have been sent from Africa ; four are from 
a collection forwarded by Mr. Claude Fuller of the Department of Agriculture of the 
Union of South Africa (Macrosiphum cassi, Phorodon violae, Capitophorus 
chrysanthemi and Forda natalensis) ; one has been sent by Mr. C. C. Gowdey from 
the British East African Protectorate which has to be placed in a new genus (Cer- 
craphis). 

Macrosiphum cissi, sp. nov. (fig. 1). 

Alate viviparous female. 

Antennae longer than body, rather dark, pale at base of third segment ; basal 
segment wider and a little longer than second ; third longer than fourth, not so 
long as sixth, with a line of 15-18 sensoria extending nearly to the end of the segment 
the distal one usually markedly separated from the others ; fourth segment a little 
longer than fifth, the latter with the usual sub-apical sensorium ; sixth with long 
flagellum and usual sensoria at apex of basal area; fourth to sixth markedly 
imbricated ; with short, simple, scattered hairs. Eyes large and dark. Frontal 
lobes small, but prominent, median ocellus prominent; head with slightly spatulate 


Fig 1. sagen eae cissi, sp. n.: A, heed af alate 2; B, cornicles and cauda of 
apterous 2; ©, cornicles and. cauda of alate 9°; D, cauda of alate 2; E, head 
and antenna of apterous eH 


hairs. Proboscis rather broad, dark at apex, reaching to or just beyond the 
second coxae, apical segment a little longer and narrower than second. Legs long 
and thin, dark at apices of femora and tibiae, tarsi dark, most of hind femora dark, 
short hairs over apex of femora and tibiae. Cornicles black, long and cylindrical, 
about as long as third antennal segment, apex reticulate, remainder imbricated. 
Cauda about one-half the length of cornicles, pale, with three pairs of long 
lateral hairs, a smaller apical pair and a median sub-apical one. Anal plate darker 
and with many hairs. Cauda not quite level with apices of cornicles. Wings 
normal. Length, 2-2-3 mm. 


66 FRED. V. THEOBALD. 


Apterous viviparous female. 

Antennae longer than body ; first to third segments pale, except apex of latter, 
remainder dark ; first segment much larger than second ; third longer than fourth, 
but shorter than sixth, with a group of 2-6 sensoria near the base ; fourth longer 
than fifth, the latter with usual sub-apical sensorium, both combined a little shorter 
than sixth to about the same length ; flagellum of sixth long; hairs normal. Head 
with well formed frontal lobes and with a small median prominence showing only 
in some specimens ; other mounted specimens show the vertex almost flat. Eyes 
large and dark. Proboscis reaching to or just beyond the second coxae ; apical 
segment longer and considerably narrower than the penultimate. Cornicles dark, 
cylindrical, somewhat expanding basally, reticulate at apex, remainder imbricated ; 
shorter than third antennal segment. Cauda pale, about half as long as cornicles 
and slightly thicker, with three hairs on each side and some apical ones ; not quite 
reaching the apices of the cornicles. Legs relatively shorter and thicker than in 
alate female; apices of femora and tibiae and tarsi darkened ; hairs moderate. 
Anal plate dusky, with rather long hairs. Hairs of body slightly spatulate. 
Length 2°2-2°5 mm. 

Nymph. 

Antennae longer than body ; darkened on fifth and sixth segments ; basal segment 
larger than second ; third longer than fourth ; fourth longer than fifth ; the sixth 
much longer than 4-++5 ; flagellum long, but relatively shorter than in mature forms ; 
a few scattered hairs. Proboscis with dark apical segment, reaching to the third 
coxae ; last two segments nearly equal. Wing pads dusky; cornicles dark ; about 
as long and as thick as third antennal segment, imbricated ; reaching past the 
cauda. Cauda triangulate, pale, with two pairs of lateral hairs. Legs with dusky 
apices to tibiae, but not to femora; latter with hairs on one side; tibiae hairy. 
Length 1°8 to 2 mm. 


Food-plant. Cissus sp. 
TRANSVAAL: Pretoria 18.x.1918. 


Described from three alate females and a number of apterous females and nymphae. 
No colour notes were sent, but the species is so well marked that I have described 
it. It comes nearest to Macrosiphum, but also approaches Myzus, especially in the 
alate stage. Mounted apterae show the vertex in different forms, some have it 
almost flat, others of typical Macrosiphum form, whilst others show a median 
prominence. In the various larval instars the differences are great; the first has 
very short thick cornicles, in later stages they increase in length. The length of 
the cornicles varies in the adults. 

The plant upon which these Aphids were found—C?ssus—is one of the Order 
Ampelideae. This genus is often merged into Vitis. None of the species recorded 
from Vtis agrees with Mr. Fuller’s specimens, and so far no Aphid has been 
recorded from Cissus. 

The Aphids recorded from Vitis are Aphis allinorsensis, Shimer ; Aphis rvparvae, 
Oestlund ; Aphis vitis, Scopoli; Macrosiphum viticola, Thomas ; Hyalopterus 
arundinis, Fabricius ; Peritymbia vitisana, Westwood ; Rhizoctonus ampelinus, 
Mokrzeki ; Schizoneura ampelorhiza, Del Guercio ; and three species of Phylloxera. 


AFRICAN APHIDIDAE.—PART IV. 67 


Rhopalosiphum carduellinum, Theobald (fig. 2). 

Bull. Ent. Res. vi, p. 112. figs. 9 and 10, (1915). 

Apterous viviparous female. 

Antennae about as long as the body, arising from prominent frontal tubercles ; 
basal segment much larger than second ; third not quite as long as the sixth, with 
17-20 small sensoria along one side; fourth slightly longer than fifth, with 3-6 
sensoria ; fifth with normal sub-apical sensorium ; sixth about as long as 4-++5, its 
basal area about one-fourth the length of the fifth ; the apices of segments 3-5 are 
darkened and there is a dark area at junction of basal area and flagellum of sixth ; 
all the segments are imbricated ; the sensoria on 3 are of varied size and shape ; 


Fig. 2. Rhopalosiphum cardueliunum, Theob., apterous female; A, 
head and antenna; B, cauda and anal plate; C, cornicles ; D, sensoria 
on 3rd antennal segment. 


those on 5 are more regular in outline. Proboscis reaching to just past the second 
pair of legs ; two bands of a few hairs on penultimate segment ; two marked sub- 
apical ones on the apical arising from clear circular areas. Legs with apices of femora 
and tibiae darkened and dark tarsi, a few hairs on apex of femora and numerous 
short ones on the tibiae. Cornicles about as long as the fourth antennal segment, 
about twice as wide and somewhat swollen on the apical half; apex with a few 
striae, remainder faintly imbricated. Cauda pale, broader than cornicles, more 
than half their length, with three long hairs on each side and a curved sub-apical 
one ; reaching just beyond the apices of cornicles. Length, 1°8-2 mm. 

Food-plant. Thistles (Carduus sp.). 

TRANSVAAL: Pretoria, 20.v.1914 (C. Fuller); Onderstepoort, 28. vii. 1914 
(G. B. Bedford). 

Type in the British Museum. 

The apterous female sent with the alatae of this insect appears to be a distinct 
species. The sensoria in the two apterous females sent by Mr. Claude Fuller appear 
so much more closely allied that I have little doubt that they are true carduellinum. 

Numbers of nymphae sent were mostly all of a uniform pale colour, evidently 
vellow or green, but a few show the tips of the wing pads dusky. One of the sensoria 
on the third antennal segment is markedly bean-shaped, and the varied size of the 
others is somewhat characteristic. ; 


Phorodon violae, sp. nov. (fig. 3). 

Apterous viviparous female. 

Yellowish green to dull pale yellow. Antennae longer than body, of same colour 
as body except extreme tip of fifth and whole of sixth segments, which are dark ; 


68 FRED. V. THEOBALD. 


basal segments (1 and 2) somewhat darker than the rest ; the basal one much larger 
than second and somewhat projecting on its inner side ; third segment longer than 
fourth, but not so long as sixth ; fourth and fifth about equal in length ; flagellum 
moderately long, about as long as third ; a few scattered hairs over the segment and 
the third with short stiff hairs arising from small projections along one side, some 
extending on to the fourth ; basal segment imbricated. Frontal lobes large, with 


Fig. 3. Phorodon violae, sp. n., apterous viviparous female: A, head of 
apterous2; B, posterior end of body and end of antenna; B. 1, cornicle; 
C, C.1, cauda. 


a marked, slightly inwardly directed, cone-like process on each side, imbricated 
and with two minute hairs. Two stiff hairs also project from the vertex. Eyes 
large, red to dark red. Proboscis pale, apex slightly darkened ; reaching to or just 
beyond the third coxae, narrow ; apical segment a little narrower and longer than 
penultimate. Legs of same colour as body, except the tarsi which are darkened ; 
very short stiff hairs on the tibiae. Cornicles of same colour as body, moderately 
long, but shorter than the fourth antennal segment, slightly swollen from the middle 
onwards ; faintly imbricated, sometimes slightly expanded at the base. Cauda pale, 
short, bluntly triangulate, spinose, with two short hairs on each side ; projecting 
well beyond the cornicles. Anal plate slightly darkened, narrow, spinose, with 
four long apical hairs and others somewhat shorter. On each side of pronotum 
a small lateral papilla and traces of four others on each side of the abdomen. The 
body is contracted at the apex. Length, 1°3-1'8 mm. 

Nymph. 

Colour similar to adult. Antennae about as long as body and of same colonir ; 
sixth segment and apex of fifth dusky ; basal segment much larger than second, 
third a little longer than fourth ; fourth and fifth about equal; sixth as long as 
or a little longer than 4++5, its basal area about half the length of the fifth. Frontal 
processes as in alate female. Eyes bright red. Wing-cases of same colour as body, 
also the legs, except apices of tibiae and the tarsi, which are smoky, with numerous 
fine short hairs. Cornicles and cauda pale, the latter bluntly triangulate and 


AFRICAN APHIDIDAE.—PART IV. 69 


extending beyond the cornicles, which are shorter than the fifth antennal segment. 
Cauda finely spinose, but showing no lateral hairs. The pale proboscis reaches 
nearly to third coxae. Length, 1°2-1°5 mm. 

Food-plant. Pansy. 

Natat: Durban, 1.x.1912. 


Described from a number of apterous females and some nymphae. Nearly all 
the females have the posterior portion of the gut, etc., evaginated, giving them a 
curious ragged appearance. The shape of the body is also marked. 

I have placed this species provisionally in the genus Phorodon on account of the 
two marked frontal processes and the swelling on the basal segment of the antennae, 
but the cornicles and cauda do not exactly fit in with that genus, taking P. hwmuli, 
Schrank, as the type. The flattened anal plate occurs, however, in some stages of 
the hop aphid. 


Capitophorus chrysanthemi, sp. nov. (fig. 4). 

Apterous viviparous female. 

Apparently a pale green or yellow species, with black eyes ; dusky apices to the 
cornicles and antennae; tarsi all dark and to some extent the apices of the tibiae ; 
hairs capitate. Antennae longer than body ; basal segment larger than second 
and somewhat angulated on the inner side ; third segment much longer than fourth 


Fig. 4. Capitophorus chrysanthemi, sp. n.: A, head of apterous 9; A. 1, 
enlarged cephalic hairs; B,analregion ; a,cauda; b,anal plate; b. 1, hairs 
of anal plate; c, dorsal process ; d, body hairs. 


but not so long as the sixth ; fourth a little longer than fifth, the latter with the 
usual sub-apical sensorium ; sixth with long flagellum, basal area about one-fourth 
of the fifth segment. Head with marked frontal lobes and a median raised process ; 
with two capitate hairs on each lobe, two in front on the median lobe, and 
two just above them, Eyes prominent, varying from black to deep red or reddish 
black. Proboscis pale, reaching to just past the second coxae; apical segment 


70 FRED. V. THEOBALD. 


narrow and acuminate, penultimate nearly as long, expanded. Legs pallid, except 
tarsi, which are dusky except at extreme base ; tibiae with moderately long fine 
hairs and a few on apex of femora. Cornicles long and very thin, about as long 
as fourth antennal segment ; pale, apex dark. Cauda pale, rather long and thick, 
bluntly elongate-triangular, with two hairs on each side, close together towards 
the apex and a median sub-apical one ; finely spinose. Anal plate slightly darker, 
spinose and with a few long hairs, broadly rounded. The thin cornicles project 
to about the level of the cauda, which is very much wider and about half to less: 
than half their length. Body hairs strongly capitate, some with fan-shaped 
extremities, whilst those on the anal plate and cauda are simple. The lateral 
view shows a small tubercle above the cauda (fig. 4, B.c.). Length, 1'8-2 mm. 

Food-plant. Chrysanthemum. 

ORANGE FREE StaTE: Bloemfontein, 18.v.1914. 

Described from several mature apterous females. It resembles a typical Myzus, 
but the median frontal projection places it in Van der Goot’s genus Capitophorus ; 
which appears to be undoubtedly well marked, but I find so many species between 
the true Myzus and typical Capitophorus that it is doubtful if this genus should 
be accepted, unless many others are to be initiated. 


Genus Cerciaphis, nov. 

Described from apterous viviparous females. 

Head flat to very slightly concave in front. Antennae shorter than body, of five 
segments, the third about as long as 4+5; the fifth longer than fourth, its 
flagellum only half the length of basal area. Kyes small. Cornicles small, cone- 
shaped. Two thick, acuminate anal cerci, about as long as fourth segment of 
antennae. Cauda small, flattened. Anal plate rounded. Proboscis rather short. 


Cerciaphis bougainvilleae, sp. nov. (fig. 5). 

Apterous viviparous female. 

Pale-coloured ; apices of antennae dusky. Eyessmallanddark. Legs, cornicles, 
anal plate, cauda and base of antennae of same colour as body. Head narrower 


Fig .5 Cerciaphis bouganvilleae, sp. n.: A, head of apterous 9 ; 
a. 1, enlarged eye; a. 2, and a. 3, sensoria; B, posterior end ; 
b. 1, cerei; ¢, cornicle; d, cauda and anal plate; C, cornicle. 


than body, which is more or less oval ; head flattened to slightly concave in front, 
integument strongly striate. Antennae shorter than body, of five segments, the 


AFRICAN APHIDIDAE,.—-PART IV. 71 


basal one larger than second, the third the longest, as long as or longer than™4-+5 ; 
the fourth shorter than fifth, with a simple round sub-apical sensorium ; the fifth 
with the flagellum short, about half the length of the basal area, which is equal 
in length to the fourth segment ; third to fifth imbricated. Cornicles cone-shaped, 
of Lachnid appearance, more or less circularly striate. Two long, pale, acuminate 
anal cerci, striate and about the length of the fourth antennal segment to a little 
longer. Anal plate pale, small, with a few longish hairs. Cauda inconspicuous. 
Legs rather short and thick. The posterior integument is striate, but not so mark- 
edly as the cephalic. The proboscis in all the mounted specimens is bent forwards! 
but appears to reach to about the second coxae. Length, 1-4—1'8 mm. 

Food-plant. Bougainvillea. 

Ucanpa : Kampala, 14.v.1919 (C. Gowdey). 

Described from a number of apterous viviparous females. It is a very marked 
insect on account of the two posterior cerci. The head and body seem almost J 
devoid of hairs, except posteriorly. The nymphae have long narrow wing-buds. 


Forda natalensis, sp.nov. (Fig. 6). 

Apterous viviparous female. 

Globular, much domed dorsally, flattened ventrally. Grey, greyish white to brown; 
antennae, proboscis and legs brown. Segmentation marked posteriorly. Antennae 
from one-fourth to one-fifth the length of body ; of five segments, basal one about 
half the length of second, third the longest, about as long as 4++5 ;}fourth shorter 


Fig. 6. Forda natalensis, sp. n.: A, head; B, cauda, C, integument 
of apterous viviparous 2; D, nymph. 


than fifth ; fifth with a short blunt nail, a large rather projecting sensorium at its 
base and traces of a secondary one, also a round sensorium at the apex of the fourth ; 
slightly hairy ; last two segments darker than remainder. Vertex flat, with a few 
short blunt hairs. Eyes very small, black. Proboscis reaching to near the third 
pair of legs or just past them, moderately broad ; apical segment a little longer 
and narrower than the penultimate, with a few short hairs. Legs rather short 
and thick, prothoracic shorter than mesothoracic, the latter shorter than the 


Te FRED. V. THEOBALD. 


metathoracic ; tibiae and tarsi with short fine hairs, a few on the femora ; femora 
of first and second pairs nearly as long as tibiae, the third a little shorter. Body 
with short fine scattered hairs, except at the apex, where they are longer. 
Length, 2-2°5 mm. 

Immature viviparous female. 

Paler, legs pale brown ; antennae dark on last two segments only. Antennae 
about one-fifth the length of the body ; basal segment shorter than second and 
very little broader ; third a little longer than second ; fourth about as long as the 
first ; fifth a little shorter than third, with a small blunt nail, with normal sensoria 
at its base ; a few scattered hairs. Proboscis reaching to or beyond the third pair 
of legs. Legs short, just projecting beyond the body, the first slightly shorter than 
second, third the longest ; tibiae and tarsi with hairs as in adult. Eyes very small. 
Body with a few rather short scattered hairs. Length, 1:5-2:0 mm. 

Food-plant. Roots of a weed and im ants nest. 

NataL: Umzinto, 23.i1v.1911. 

Described from several apterous viviparous females and some immature forms. 
It somewhat resembles Forda rotunda, Theobald, but can at once be distinguished 
by the long second antennal segment and the short fourth segment, also by the 
flat vertex and different posterior structure. The eyes also do not project in the 
same way, and the body is clothed with scattered fine short hairs. 

Some specimens show a partial constriction on the third antennal segment, 
but I could find none in which it was complete, so that they are only of five segments. 


Species not previously recorded or from new Localities. 


Macrosiphum dahliafolii, Theobald, Bull. Ent. Res. vu, p. 273, 1918. 
Ucanpa: Kampala, 2.x11.1917 (Gowdey) ; on thistles (Carduus). 


Macrosiphum sonchi, Linnaeus. Syst. Nat. u, p. 735. 

UGanDA: Kampala, 2.xii.1917 (Gowdey) ; on thistles (Carduus). 

One apterous 2, which agrees with British specimens. The darkening of the 
antennae around the sensoria of the third antennal segment is very marked. 


~I 
Ww 


ANOPHELES BREEDING AMONG WATER LETTUCE—A NEW HABITAT. 


By James ZETEK, B.A., 


Entomologist, Ancon, Panama Canal Zone. 


With the filling up of Gatun and Mira Flores Lakes, water lettuce (Pistia stratiotes, 
Linné) and water hyacinth (Hichornia crassipes, Solms.) rapidly increased in numbers 
until large areas became completely covered by them. Masses of these plants would 
become detached and float about in these lakes. It was found necessary to destroy 
these “floating islands,” not because they were serious mosquito habitats, but 
because they interfered with navigation and the operation of the spillways. The 
water lettuce is the habitat of the very specialised larvae of Mansonia, of which 
titillans is the commonest member. Knab (1913) and Busck (comments appended 
to Mr. Knab’s paper) refer to the changes in the mosquito fauna brought about by 
the rapid increase in Pistia. Both Mansonia titillans. Walker, and Aédomyia 
squamipennis (Lynch-Arrib.) Theo., have been collected by me in very large numbers 
since 1912, and to-day Mansonia is the dominant species caught in the army barracks 
on the west side of the canal. Knab expressed the opinion that perhaps measures 
would have to be taken to destroy the Pistia habitats ; now that we have found 
Anopheles larvae in them, this prophesy comes closer home. 

The only question involved is how much of a menace are these floating islands 
to our towns. It is my belief they are a menace to the towns on the west side 
of the canal, and if allowed to float toward Paraiso and Pedro Miguel, or close to 
Gatun, would be a source of danger there. It has been shown conclusively that our 
common malarial Anopheles do fly more than a mile (Zetek, 1915) ; in fact, it appears 
that they will fly as far as they must in order to get food. Should Las Cascades 
become a negro settlement, the dangers from malaria would be increased. I doubt 
if we can depend upon screening and mosquito-catching indoors to control the 
Anopheles and malaria at such a settlement. At any rate, the idea seems to be 
getting firmly established that the sanitation of the Panama Canal Zone is so efficient 
that there is virtually little danger from the Anopheles, and as a result our people 
are becoming somewhat negligent. It will depend mainly upon the cumulative 
evidence gathered by the District Sanitary Inspectors, whether or not these floating 
islands will need to be controlled. 

That this evidence may be available and accurate, it is very necessary that the 
sanitary Inspectors make it their practise to take ample field notes. These should 
contain as complete a statement of the particular habitat, date and locality added 
thereto, and specimens of the larvae should be sent to the laboratory for the accurate 
determination of the species. At the close of each month a general summary of 
these notes should be made, a copy of which should be sent to the entomologist. 
It is a regrettable fact that much valuable information is lost or buried in the heads 
of observers. 

In 1918 I found Anopheles larvae among water lettuce at Juan Mina, a citrus 
plantation about five miles up the old Chagres River. On 21st November 1919, 
Messrs. Picket and Tolar, sanitary inspectors at Pedro Miguel, D>. D. P. Curry, 

(681) F 


74. JAMES ZETEK. 


Assistant Chief Health Officer, and I, made an inspection of floating islands at 
Gamboa, C. Z., both in the canal prism and across in the bayous ; we went as far 
as Empire. Mr. Pickett previously reported the presence of Anopheles larvae 
in these islands at this place, and that they were going southward toward his station. 
Our joint inspection revealed larvae of Anopheles albimanus, Wied., and 
A. tarsimaculatus, Goeldi, to be plentiful among the leaves of the water lettuce. 
An unidentified Culex was also present. 

This particular habitat is very favourable to the Anopheles. The larvae are 
afforded much protection from the schools of young fish that usually feed upon 
insect larvae. They are also protected to some extent from predaceous insect 
larvae, such as those of dragon-flies. Shade is another important factor in this 
habitat, and this means a more even temperature, and an almost total absence of 
the direct hot rays of the sun. Wave action does not disturb these larvae very 
much ; but the most important advantage to them is the exceptionally favourable 
presence of oxygen, and this certainly must be effective in prolonging the life of 
the larvae and pupae. This factor is very easily explained. The Pistia, being 
a green plant, gives off oxygen. Its leaves are at or just above or below the surface 
of the water. Therefore this oxygen enriches the local atmosphere available to 
the larvae, and some of it is undoubtedly taken up by the water and made use of 
by the larvae through their cutaneous respiration. 


During the summer of 1911, while engaged by Dr. 8. A. Forbes to study the 
Anopheles at Havana, Illinois, a similar condition was found to exist, excepting 
that in place of Prstia the plant in question was the tiny Lemna. This plant covered 
the surface of the water in the bayous, ponds and quiet recesses of the rivers. The 
Chatagua grounds at Havana were invaded each year by Anopheles, but their source 
was not known until we found them in and among the floating islands of this Lemna. 
These islands broke away from the almost continuous mass across the river from 
Havana and by current and wind were swept across this deep body of water and 
came to rest along the shores of the Chatagua grounds ; here the larvae developed, 
pupae formed and adults emerged in rapid succession. 

There is one important aspect of this sort of habitat which must not be over- 
looked. It is that wind and current detach portions of this floating vegetation 
and sweep it across deep bodies of water to new localities, where were it not for this 
exceptionally good vehicle these larvae would be unable to cross the barrier of deep 
water. 

Searching through the literature available at our laboratory, I find that Ingram 
and Macfie (1917) found larvae of Anopheles costalis, Lw., and A. marshalli, Theo., 
in exactly the same sort of habitat, composed of the same species of Pistia, at 
Christiansborg, near Accra, West Africa. Associated with them were: Aédomyra 
africana, Nev.-Lem., Mimomyia splendens, Theo., Culex quasigelidus, Theo., and 
Mansonwoides africanus, Theo. 

In a previous paper, in which he discusses the limitations of kerosene as a larvicide, 
Macfie (1917) states (p. 278) that Mansoniordes africanus flies over one mile and that 
its breeding places can be located with fair accuracy, inasmuch as they are composed 
of Pistia. On page 278 he states it is useless to hope for a total abolition of such 
mosquitos by ordinary oiling, and that a layer of kerosene (p. 294), no matter 


ANOPHELES BREEDING AMONG WATER LETTUCE. 75 


how thick, applied to the surface of the water in which larvae of M. africanus live 
is absolutely innocuous. He is right, and this is corroborated by our observations 
upon Mansonia titillans, which is our counterpart to his M. africanus. But nowhere 
in this article, nor in the preceding one, does he touch upon the control of the Ano- 
pheles which live in this same habitat, nor does he refer to the use of emulsions which 
mix with the water and kill by contact, or the destruction of the Pistia by means 
of sodium arsenite or their removal by boats. 

The logical control is to destroy the habitat. This may be costly and prolonged, 
and perhaps prohibitive where large numbers of bayous, ponds, cut-ofis, etc., exist. 
If Anopheles alone are to be controlled, spraying with a phenol-resin soap emulsion 
every six days will probably be all that is necessary. It may even be effective 
against the Mansonia. 

Macfie (1917) goes into great detail in his discussion of the favourable oxygen 
factor in this Pestia habitat and relates experiments made with certain mosquito 
larvae which corroborate his contentions. That this exchange of gases is a favour- 
able one is quite certain. Our common Anopheles have been known to stay on the 
bottom for five minutes ; if through the presence of green plants they have a more 
favourable habitat, then it follows that not only may cutaneous respiration be much 
more prolonged, but the general death-rate among the larvae may be greatly reduced. 
This death-rate is reduced also because the leaves of the plants protect the larvae 
from the numerous swimming enemies in the water, and from the hot, direct rays 
of the tropical sun. 


References. 


Incram A. & Macriz J. W.S. (1917). The Early Stages of Certain West African 
Mosquitos.—Bull. Ent. Research, vii, pt. 2. 

Kwap, Frederic (1913). Changes in the Mosquito Fauna of Panama.—Proc. Ent. 
Soc. Washington, xv, pp. 41-43. 

Macrie J. W.S. (1917). The Limitations of Kerosene as a Larvicide, with some 
Observations on the Cutaneous Respiration of Mosquito Larvae—Bull. Ent. 
Research, vii, pt. 3. 

ZETEK, J. (1915). Behavior of Anopheles albimanus, Wied., and tarsimaculata, 
Goeldi.—Ann. Ent. Soc. America, viii, pp. 221-271. 


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LIST OF THE ANOPHELINES OF THE MALAY ARCHIPELAGO WITH 
SPECIAL REFERENCE TO ADULTS AND LARVAE OF NEW OR 
INCOMPLETELY DESCRIBED SPECIES OR VARIETIES. 


By N. H. SwELiteNnGREBEL, Ph.D. and J. M. H. SwWELLENGREBEL DE GRAAF. 


The following species and varieties have been found in the Malay Archipelago* :— 
Myzomyia: ludlowi, rossii, vaga (indefinita), flava, minima, minima var. aconita. 
Neomyzomyia : leucosphyra, punctulata var. tesselata. 

Cellaa : kochi. 


Myzorhynchus : sinensis, sinensis var. vanus and var. separatus, barbirostris, 

barbirostris var. pallidus, albotaeniatus, mauritianus, umbrosus, gigas. 

Nyssorhynchus: fuliginosus, maculatus, schiiffneri, karwari, jamesi, annulipes 

var. moluccensis. 

Stethomyia : aitkenii, aitkenii var. insulae-florum and var. papuae. 

Comparison with the neighbourmg Malay Peninsula shows the Anopheline 
fauna to be almost identical in the two countries. The exceptions are :— 
M. aurirostris, P. watson, M. sinensis var. peditaeniatus, M. albotaeniatus var. 
montanus, M. pseudumbrosus, M. hunterr and L. asiatica, which are not found 
in the Archipelago; whereas the Peninsula lacks M. mauritianus, M. gigas, 
M. barbirostris var. pallidus, S. aitkenw var. insulae-florum and var. papuae, and 
N. annulipes var. moluccensis. 


I.—DeEscRIPTION OF NEW SPECIES AND VARIETIES. 
1. Myzomyia immaculata, Theo. 
Myzomyia flava, Swellengrebel (1917).T 
A yellow mosquito, with light orange-coloured unbanded legs, creamy unbanded 
palpi, brownish yellow proboscis and creamy wing scales. Wings unspotted. 


2. Head scaled like M. rossw, on the occiput broadly expanding upright forked 
scales, all scales creamy. Palpi unbanded, creamy. Proboscis brownish yellow, 
labella yellow. Thorax: prothoracic lobes with brownish yellow hairs. Mesonotum 
with hair-like scales, narrow curved ones in front and a few at the sides, all scales 
creamy. Scutellum with dark yellow hairs and creamy hair-like scales. Halteres 


* Contrary to the present custom we adhere (with a few exceptions) to Theobald’s 
nomenclature, not because we attach to his ‘genera’? any intrinsic value as indicating 
a reaily existing relationship, but because the division of the Anophelines into genera 
is a practical measure, expressing the idea present in the mind of every one who studies 
these insects in nature, that, e.g., M. sinensis is quite distinct from M. aconita, but closely 
allied to M. barbirostris, and that ‘8. aitkentt i issomething apart from all other Anophelines. 
By calling all these species ** Anopheles’ one simply causes confusion and not simpliliea- 
tion. 


{ [On a recent visit to England Dr. Swellengrebel presented to the British Museum a 
6 and 2 of his Myzomyia flava. On comparing the latter with the type of Theobald’s 
A. immaculatus both he and Mr. F. W. Edwards concluded that they were conspecific. 
The older name immaculatus must therefore be used for this interesting and little 
known form.—EbD. | 


78 N. H. SWELLENGREBEL AND J. M. H. SWELLENGREBEL DE GRAAF. 


creamy, light brown at the curve. Abdomen with golden hairs, narrow curved 
golden scales on genital lobes. Legs unbanded, light orange. Ungues equal and 
simple. Wangs unspotted; scales narrow, creamy or light yellow. Base of Ist 
submarginal cell nearer the wing base than that of 2nd posterior ; the cell is longer 
than its stem, 2nd posterior cell shorter than its stem. Supernumerary and 
anterior cross-veins meeting, the posterior one more than twice its length from the 
latter ; 3rd longitudinal vein not ending at the meeting of the supernumerary and 
anterior cross-veins. Length, 4-5 mm. 

§. Like 9. Club-shaped end of palpi yellow, owing to the presence of long 
yellow hairs mixed with the creamy scales ; a dark narrow band caused by partial 
desquamation. Narrow-curved creamy scales on the two apical abdominal seg- 
ments. On the wing, anterior cross-vein situated basally from the supernumerary. 
Larva unknown. 

JAVA: Soerabaia. SumaTrRA: Mandailing. 

Rare. Described from 3 99 and 3 J taken at Soerabaia, June 1917. Easily 
distinguished from other unspotted Anophelines by the long palps (M. brevipalpis) 
and the broadly expanding upright forked scales on the head (S. aitkeni). The 
species may perhaps be an albinoid variety of M. vaga (indefimita), as this species 
sometimes shows a marked decrease of the black portions on the wings and palpi. 


2. Nyssorhynchus annulipes var. moluccensis, Swell. (1920). 

A rather large brown mosquito, with spotted legs, 4-5 light bands on palpi, 
the apex of the 2nd joint being black; proboscis dark, except labella; narrow 
curved scales on the whole dorsal surface of the mesonotum ; wings much spotted, 
resembling those of NV. punctulata var. tesselata. 

2. Head scaled as in other Nyssorhyncht. Antennae light. brown, basal segment 
with small white scales, verticils and tomentum golden. Proboscis dark brown, 


Fig. 1. Palpi of Nyssorhynchus annulipes var. 
moluccensis 9, three variations. 


labella lighter. Palpi (fig. 1): 1st joint black with creamy apex and sometimes a 
dorsal yellow spot in the middle; 2nd joint black, with a broad white band on 
apical half and a narrow yellow one at the apex, sometimes a yellow dorsal spot 
in the middle of the black basal portion, the sub-apical white band sometimes narrow 
or even absent; penultimate and terminal joints with a narrow black basal band, 
then a narrow yellow band, the remaining portion creamy. Thorax: prothoracic 
lobes blackish brown, with black chaetae and a tuft of black scales in front. Meso- 
notum with a dark spot to the right and left at one-third from the anterior border 
and one in front of scutellum, covered all over with narrow curved white scales, 


. 


LIST OF THE ANOPHELINES OF THE MALAY ARCHIPELAGO, T9 


which also are present on scutellum. Pleurae dark, with a few narrow curved 
white scales. Metanotum light brown, with a dark median line. Halteres white, 
with the knob dark at the apex. Abdomen dark brown; on apex of 6th and 7th 
segments narrow curved golden scales, on 8th black ones likewise. Legs dark brown 
with white spots, many of them forming incomplete rings; 1st tarsal joint banded 
apically, 2nd—-4th at both ends (except in hind legs, which have an apical band only) ; 
5th banded at both ends in front legs, unbanded in the others, in front and mid-legs 
this joint is white on one side. A sub-apical elongated light spot on the tibiae of 
front legs. Coxae yellowish brown, with narrow white scales. Wings (fig. 2): 


ee — ee 
~ ae a ce, ot 
Co aa sae =" 


leg tea a == i? 


Fig. 2. Wing of N. annulipes var. moluccensis , 2. 


on the costa from base to apex 3 short and 4 long black spots and a short one between 
the 1st and 2nd long spots, these last three extending on to the sub-costa. The 
1st-4th long costal spots extend on to the Ist longitudinal in the form of 2, 3-4, 34, 
small spots and one long one; moreover a small spot below the interval of the Ist and. 
2nd and the 2nd and 3rd costal long spots, and 1-3 at the base. Other small spots 
are present as follows :—3-5 on upper branch, 5-7 on lower branch, 24 on stem of 
2nd vein; 5-6 on 3rd vein; 2 apically, one longer basally on upper branch, 2-3 on 
lower branch of 4th vein, on its stem below the cross-veins a long one, above them 
2 long or 1 long and 2 short ones; 4 on the upper branch and lower one, 3 on the 
stem of the 5th vein; 5-6 on the 6th vein. Fringe dark with bright incisions at 
the tips of the veins and upward from the 3rd longitudinal vein, except a narrow 
black spot between the branches of the 2nd vein. Length of 1st submarginal cell 
more than twice that of stem, its base situated basally from that of 2nd posterior 
cell. Supernumerary and anterior cross-veins almost meeting, posterior one twice 
its length from anterior. 
Length, 4—5°5 mm. 


3. Palpi: 1st joint black, mixed with white on one side; articulation with 2nd 
joint light, owing to absence of scales ; 2nd joint with one side of basal half and top 
white, the latter white portion fringed with yellow. Club-shaped end white, black 
on one side, with basal and median black ring, separated from the white by a yellow 
margin. Length 5-6 mm. 


Mouuccas. New GUINEA. 
Described from 592° and 34. 

Larva (fig. 3). Length 5 mm. Colour greenish brown or dark greyish-green ; 
head brown, with dark bands and spots; white spots on thorax in front and on 
abdominal segments iii and vii or ili and vi-viii. Antennae without branched hair. 
Median clypeal hairs far apart, with a few short hairs ; external clypeal hairs often 
nearly as long as medians, with long hairs ; posterior clypeal hairs as far apart as 
medians, reaching to the base of the latter, unbranched or with 2-3 branches. 


80 N. H. SWELLENGREBEL AND J. M. H. SWELLENGREBEL DE GRAAF. 


External occipital hairs with 4 branches, internal ones with 3 branches or simple. 
A pair of small fans* on thorax, with 10-14 leaflets, sometimes pigmented, no 
serrate margin. Small fans on abdominal segment i, with 8-16 leaflets, usually 
serrate, not pigmented. Complete fans on segments li-vii, with about 18 leaflets, 
pigmented, serrate near the tip, the latter long and pointed, sometimes ending 
in a true filament. Shoulder hairs f with thick stem and numerous branches ; median 
and internal one with stout roots which often unite into one. 


Fig. 3. Larva of NV. annulipes var. moluccensis: a, b, two types of 
clypeal hairs; c, occipital hairs; d, shoulder hairs (right); e, f, 
leaflets. 
Breeds in all kinds of water, brackish or salt and fresh, running and stagnant, 
clean and dirty ; also in artificial collections of water (coconut-shells, water in 
native boats). 


* Fan = stellate tuft. 
t+ Shoulder hairs = anterior thoracic hairs. 


LIST OF THE ANOPHELINES OF THE MALAY ARCHIPELAGO. 81 
' 


This variety much resembles N. annulipes, Walker, as described by Theobald, but 
the female palpi are different in the latter, the apical half of the 2nd joint being 
all white. Also, there is a marked likeness to N. punctulata, Dén., but the proboscis 
of this species has a light apical half and the black apical ring of the 2nd palpal 
joint is much narrower. 


3. Stethomyia aitkenii var. insulae-florum, Swell. (1920). 

The imago is not to be distinguished from S. aatkeni. 

Larva (fig.4). Length5mm. Colour dark green. Antenna with small branched 
hair. Median clypeal hairs close together, simple, nude; external clypeal hairs 
simple, nude, length 4 of median ones; posterior clypeal hairs far apart, short, 
with 3-6 branches ; occipital hairs with 3-6 branches. Shoulder hairs with only 
the median one inserted on a strong root, carrying about 11 branches, the internal 
one with about 8. Small fans on thorax and abdominal segment 1, carrying 13-15 


Ve 


Fig. 4. Larva of Stethomyia aitkenit var. insulae-florum : a, clypeus ; 
b, antenna; ¢c, shoulder hairs (left); d, leaflets. 


= 


non-serrate leaflets ; segments ii-vii with complete fans, the leaflets very indistinctly 
serrate, ending in a long point, no filaments. 

Breeds in running water in the weeds along the edges, in hilly regions. With the 
variety papuae it is the only representative of this species in the Australian region 
of the Archipelago. Only once has it been found in Java (Island of Noesa 
Kembangan on the South Coast). 

The larva is somewhat like that of S. culiciformis, James, but the club-shaped 
antennal hair is wanting and the balancing hairs on abdominal segments 1-11 are 
of the usual type and number. 


4. Stethomyia aitkenii var. papuae, Swell. (1920). 

Although no adults could be bred from this larval variety, we feel sure that it 
likewise belongs to this species. It much resembles the larvae of the preceding 
variety, but is separated from it by the fact of the internal shoulder hairs being 
very small and carrying two branches only. The leaflets of all the fans show no signs 
of serration. 


82 N. H. SWELLENGREBEL AND J. M. H. SWELLENGREBEL DE GRAAF. 


The larvae described here, were full-grown, the pupal hairs already showing 
through the integument of the 1st abdominal segment. 

Breeds in the same places as the foregoing variety, but was only found in New 
Guinea (Kokas, Kaimana, May 1919). 


4. Myzorhynchus barbirostris var. pallidus, nov. 

Adult like M. barbirostris. 

The relation of this variety to its type is the same as that of M. peditaeniatus, 
Leic., to M. sinensis, viz., the larvae differ from the typical ones by the external 
clypeal hairs bearing a smaller number of branches (11-22 against 60 or more in 
the type) (fig. 5). 


| 


Fig. 5. Larva of Myzorhynchus barbirostris 
var. pallidus, right half.of clypeus. 


Breeds in slowly running water, in springs in the jungle and in rice-fields, but 
always in hilly country. It is rare in Java and Sumatra; in the Australian portion 
of the Archipelago it is as common as the typical form. 


II. LARVAE OF KNOWN SPECIES NOT YET OR INCOMPLETELY DESCRIBED. 


1. Myzorhynchus albotaeniatus (fig. 6). 

Length 5-5°5 mm. Colour dark rusty brown, lighter on dorsum of 3rd abdominal 
segment, less conspicuously so on the 6th, the 4th almost black. Antennae more 
stumpy than in M. sinensis, length : breadth: :1:4 (1:5 in snensis) ; a branched 
hair on antennae, its stem half the length of the hair (in sen msis 4-1), the branches 
emitted nearly at right angles (in sinensis at very sharp angles). Median clypeal 
pairs simple, nude, the distance between them equal to that separating them from 
the external clypeal hairs (close together in sinensis) ; external clypeal hairs with 
18-24 terminal branches; posterior clypeal hairs short, with 3 branches ; occipital 
hairs with 6-7 branches. Internal shoulder hair very small, with 2-3 short apical 
branches. Small fans with 12-16 non-serrate leaflets on thorax and abdominal 
segments i-ii; complete fans on segments iii-vil ; leaflets as in M. sinensis. 


LIST OF THE ANOPHELINES OF TilF MALAY ARCHIPELAGO. 83 


Breeds in slowly running jungle streams with scanty vegetation but much vege- 
table detritus. Java and Sumatra. 


6b 


“ 


Fig. 6. Larva of Myzorhynchus albotaeniatus: a, clypeus; 5b, antenna ; 

c, occipital hairs’; d, shoulder hairs (right); e, variation of inner shoulder hair ; 

f, leaflet, thoracic fan; g, leaflet, fan on abdominal segment i; h, leaflet, 
segmentiv ; j, leaflet, segment v. 


2. Neomyzomyia leucosphyra (fig. 7). 

As our own observations differ from those of Stanton (1915) the larva is redescribed 
here. | 

Length 4:°5-5°5 mm. Colour light yellowish brown or brownish grey. Antennae 
without branched hair. Median clypeal hairs long and slender, simple, sometimes 
carrying a few minute hairs; external clypeal hairs nude and simple; posterior 
ones short, nude, simple; occipital hairs with 2-3 branches. Shoulder hairs 
usually pigmented, median and internal ones with strong stems, numerous branches 
and stout roots, often uniting into one. Small fans on thorax, with 10-11 non- 
serrate leaflets. On abdominal segment i very small fans with 5-7 hair-like leaflets ; 


84 N. H. SWELLENGRELBEL AND J. M. H. SWELLENGREBEL DE GRAAF. 


on segment ii small fans with 8-11 very narrow leaflets ; on segments iii-vii complete 
fans with about 18 leaflets each, showing 2-3 indentations near the apex, the latter 
pointed or rounded, no filament. The leaflets are pigmented throughout. 


Breeds in stagnant shady freshwater pools. 


e 


Fig. 7. Larva of Neomyzomyia leucosphyra: a, clypeus; b, shoulder 

hairs (left) ; ¢, occipital hairs ; d, leaflet of thoracic fan ; e, leaflet of fan 

on abd. segment i; f, leaflet on segment ii; g, leaflet on segment iii ; 
h, leaflet on segment v. 


3. Myzorhynchus umbrosus. 

Addition to Stanton’s description. Another larval type was usually met with 
in salt or brackish water near the coast, with median clypeal hairs bearing numerous 
short hairs on the apical half. The adult emerging from this was of the usual 
umbrosus type. 


LIST OF THE ANOPHELINES OF TRE MALAY ARCHIPELAGO, 85 


4. Myzomyia minima var. aconita (fig. 8). 

Addition to Stanton’s description : In running water a larval type was common 
with completely nude median and simple long posterior clypeal hairs. The adult 
emerging from this larval variety showed the usual characteristics of aconita. 
The breeding places of the two larval types differ somewhat: out of 644 typical 
larvae 42 per cent. were found in rice fields, 11 per cent. in running water, 31 per 
cent. in fish-ponds, 15 per cent. in marshes and | per cent. in dirty stagnant water ; 
the percentages of 562 specimens of the larval variety found in the same breeding- 
places were 1°7, 88, 10, 0°3 percent. and mil. Consequently the latter is more adapted 
to life in running water. 


Rents 


eae ie 


Fig. 8. Myzomyia minima var. aconita, larval 
variety : a, clypeus ; D, leaflet of fan on abdominal 
segment Vv. 


It is to be noted that the larva of the typical MZ. minima (fig. 9) is almost identical 
with the larval variety of aconita, the only difference we could detect existing in the 
length of the filaments of the leaflets which is }—3 of that of the whole leaflet (in the 
larval variety of aconita this relation is usually 35-3). 


5. Differences between the Larvae of Nyssorhynchus punctuilata var. tesselata and 
Cellia kochi. 

Stanton enumerates them as follows: In N. punctulata there are no fans on 
abdominal segments 1 and 1, whereas in C. kochi they are present on the latter. 
We noted in both species on segments 1 and ii small fans with very narrow or 
hair-like leaflets ; in C. kochi on segment i sometimes a cockade only. Consequently 
in the Malay Archipelago the fans cannot be used to distinguish these larvae. We 
use the following characters: In N. punctulata: (a) Antennae usually much pig- 
mented ; (b) occipital hairs short with 3 or more branches; (c) internal shoulder 
hairs short, usually with no more than 3 apical branches. 

In C. kochi: (a) Antennae light ; (6) occipital hairs longer, unbranched or bifur- 
cated ; (c) internal shoulder hairs long, with numerous long branches inserted at 
intervals along the stem. 


86 N. H. SWELLENGREBEL AND J. M. H. SWELLENGREBEL DE GRAAF. 


In the Australian region of the Malay Archipelago, a larval variety of N. punctulata — 
is commonly met with, bearing fans with non-serrate leaflets, no fans but cockades 
on the 2nd abdominal segment, and a small fan with hair-like leaflets on the Ist. 


2 


d te 


Fig. 9. Larva of Myzomyia minima: a, clypeus ; 

b, occipital hairs (left); ¢, leaflet from fan on 

abdominal segmenti; d, leaflet on segment ii ; 
e, leaflet on segment vi. 


6. Difference between the Larvae of Myzorhynchus sinensis and M. barbirostris. 

In addition to Stanton’s differential characters we noted the following, which 
however cannot be used if the larvae do not show white spots on the dorsum of the 
abdomen. But if these are present, they show in M. sinensis on abdominal seg- 
ments ii, v, vill, in M. barbirostris on the segments iii and vi-vui but never on v. 
N. annulvpes var. moluccensis has abdominal spots like M. barbirostrvs. 


7. Abnormal Larvae of Myzorhynchus sinensis and M. barbirostris (fig. 10). 


In both species one of the median clypeal hairs may be placed in close proximity 
to the neighbouring external clypeal hair, at the same time exhibiting on a shortened 


LIST OF THE ANOPHELINES OF THE MALAY ARCHIPELAGO. 87 


stem a number of long apical branches (10 or more), somewhat resembling the ex- 
ternal clypeal hairs of M. uwmbrosus. 


Fig. 10. Abnormal clypeus of larva of 
Myzorhynchus sinensis. 


8. Variations in the larvae of Myzomyia ludiowi and M. rossii. 


As yet we have not succeeded in differentiating these species ; in both we have 
noted the following variations of the mature forms: (a) The posterior clypeal hairs 
are of normal length and bifurcated, or much shorter and trifurcated ; (6) the 
median clypeal hairs are bifurcated ; (c) all clypeal hairs bear 2-5 long apical branches. 

These variations are infinitely more common in M. rossi than in M. ludlowt. 


Sometimes we found M. rossit emerging from larvae resembling those of M. vaga 
(endefinita). 


9. Variations of the Larvae of Myzomyia vaga (indefinita). 


We never found M. vaga with larvae like M. ludlowi, they always adhere to the 
type first described by Strickland (1915), viz., with the short external and posterior 
clypeal hairs, the latter placed at a short distance behind the median clypeal hairs 
and closer together than these. As variations we have noted the bifurcation of the 
median and posterior clypeal hairs, the latter being placed farther backwards ; also 
they may be placed close together, the one a little in front of the other. 


10. Nyssorhynchus schiffneri (fig. 11). 


The larva was described by Mangkoe Winoto (1918). They are difficult to distin- 
guish from those of N. fuliginosus. The following are the characters which separate 
them : (a) the median clypeal hairs bear more and longer branches ; (}) the posterior 
clypeal hairs are unbranched or bifid ; (c) the stems and roots are less thick than in 
N. fuliginosus ; (d) the filaments of the leaflets on the fans are somewhat shorter 
than in the same organs of N. fuliginosus. 


88 N. H. SWELLENGREBEL AND J. M. H. SWELLENGREEEL DE GRAAF, 


Fig. 11. Larva of Nyssorhynchus schuffneri: a, clypeus ; b, leaflet of fan on 
abdominal segment v. 


Ill. QUESTIONS CONCERNING THE VALIDITY, DEFINITION AND NOMENCLATURE 


OF SOME SPECIES. 


1. Nyssorhynchus fuliginosus var. nivipes. 

This variety is characterised by the followmg points: ‘~) the white area on 2nd 
joint of the hind tarsus is very extensive, 3-3 of the jot. —_‘b) wing much lighter, 
with smaller spots, notably on 3rd vein and upper branch or 5th. The same wing 
markings are present in the male of the typical N. fuliginosus, consequently they 
cannot be used to separate the males of the type and the variety. 


2, Myzomyia minima, and var. aconita. 

Christophers (1916) separates these two forms on the grounds that the type (qa) 
has the proboscis in the 9 dark, or with only a light ventral spot near the apex ; 
(b) lacks a light frmge spot at the apex of the 6th vein ; (c) has the apical half of the 
6th vein completely dark; (d) has a long black spot on the base of the 3rd vein. 
To this we may add that (e) the type shows a broad black band on the middle of 
the light apex of the palps (in aconita this band is narrow). The two forms seem 
well separated by these characters, but we have met with otherwise typical aconita 
exhibiting the pomts (b) and (c), or (a), or (b) (c) (e). Moreover, we have found 
forms with the points (b) and (c) complete and (qa), (d), (e) intermediate ; 7.e., with 
a large light patch on the proboscis but ventrally only, a small black patch on the 
base of the 3rd vein and a band on the apex of the palps in breadth intermediate 
between that of manima and aconita. On the other hand, we have observed speci- 
mens bearing a spot on the base of the 3rd vein at least equalling in length 4 of the 
stem of the lst submarginal cell, but with all the other minima characters present 
likewise. 


LIST OF THE ANOPHELINES OF THE MALAY ARCHIPELAGO. 89 


As we do not think it advisable to throw these two forms together, we propose 
a purely artificial but easily observable distinction, viz. :— 
(a2) No spot on base of 3rd vein... M. aconita, Don. 
(6) A small spot present (length about } sf iifts Ist sihwiscomnal cell), 
M. aconita var. cohaesa, Don. 
(c) A long spot present (from half as long to as long as this cell)... minima, Theo. 


3. Myzomyia vaga (indefinita). 

The species known under the name of M. indefinita, Ludl. (1904) is separated 
from M. rossit by (a) a narrow black band at the base of the light apex of the female 
palpi ; (b) a light spot at the apex of the proboscis in the female, behind the labella ; 
(c) a lighter hee (d) its larger size ; (e) the larva. The points (a), (c) and (d) 
are subject to great variation, (b) and (e) being constant and reliable ; as (5) is not 
present in the males and (a) only refers to the females, the males cannot be dis- 
tinguished with certainty. 

In 1902 Dénitz described under the name A. vagus a mosquito distinct from 
although resembling M. rossu. The proboscis of the female is described as being 
black, with light apex and whitish labella. The palpi of the female are black, 
the terminal joints white, the penultimate with a broad black band around the base. 
A distinct variety from Celebes has the penultimate joint of the female palpus 
white only at the apex. As this is the condition actually existing in M. rossii we 
infer that the specimen on which the species vagus was founded had a narrower 
black palpal band. The species was described from a female caught at Fort de 
Koch (Sumatra) and we know that no M. rossw has ever been found there. From 
the description of the proboscis and palps of the female (in connection with the 
description of the variety from Celebes) and the origin of the type we believe that 
there can hardly be any doubt that Dénitz’s vagus and the species now called 
A. indefinitus are identical. 

Christophers (1916) suggests that Giles’ M. rossi might be the one with the 
broad white apex of the palps. As Giles (1900) states that the apical half of the 
end joint (2.e., of the two terminal joints taken together) is white in the female, 
we believe this supposition to be not well founded. On the other hand we believe 
him to be quite right in suggesting that Ludlow’s indefinita is indeed M. rossii, 
Giles, as she distinctly states (1904) that the palpal markings and general colour 
of her new species are like those of M. ludlowz. 


4. Difference between Myzomyia ludiowi and M. rossii. 

The following wing marking is very constant and reliable: on the upper branch 
of the 5th vein, below the cross-veins, both species show two spots, which however 
in M. ross (and M. vaga) are short, whereas in M. ludlowi they are much longer. 
An extensive biometrical research has convinced us that exceptions to this rule are 
rare (Swellengrebel 1916, Mangkoe Winoto 1918). 


5. Neomyzomyia punctulata and N. tesselata (A. deceptor, Don.). 

One of us has shown already that the characters differentiating these two species, 
so far as they are based on the wing markings, cannot be accepted as valid. Con- 
sequently the species are separated only by the palpal markings: in N. tesselata 

(681) G 


90) N. H. SWELLENGREBEL AND J. M. H. SWELLENGREBEL DE GRAAF. 


the second palpal joint of the female is white on the apical half, in N. punctulata it 
bears a black sub-apical ring, separated however by a narrow light ring from the 
basal black ring of the 3rd joint. We do not consider this difference of sufficient 
value to justify a new species and consequently we rank J. tesselata as a variety 
of N. punctulata. 

Donitz believed the var. tesselata to be peculiar to the western regions of the 
Malay Archipelago only, but we have found it as far eastward as Ceram (Moluccas). 
Dénitz’s typical punctulata we have never met with. The eastern punctulata var. 
tesselata differs however from the common western type, (a) by the larvae (p. 7.), 
(b) by the light portion of the proboscis being distinctly diminished in size. 


6. Nyssorhynchus schuffneri. 

Christophers (1916) seems to suggest that this species resembles N. fuliginosus 
var. adie, James. It is said to differ from it by the very short ultimate joint of the 
female palpus. We found the length of this jomt to be 11 per cent. of the palpal 
length, the palpal index being 0°6, consequently it is an orthodactylous mosquito. 
But there are other, more striking differences. The palps have 3 white bands only 
(4 in adier), with a long white tip resembling M. vaga. The wing, which in the var. . 
adiei is like that in N. fuliginosus (judging by James’ description), is much lighter 
in schiiffnert ; the black spots on the costa are rather narrow, on 2nd longitudinal 
_ vein one small spot under the 3rd costal spot (reckoned from the apex). Stem 
of 4th longitudinal vein nearly all yellow, so is the 3rd vein (in adie it is black, 
except for a few minute light spots). 


IV. GEOGRAPHICAL DISTRIBUTION. 


A considerable portion of the Malay Archipelago has been searched for Anophelines, 
and although much remains to be done, we wish to draw attention to the following 
striking features in the geographical distribution of certain species and varieties. 


1. Nyssorhynchus annulipes var. moluccensis. 

Wherever it occurs this is a very common species, breeding everywhere, and so 
it is difficult to overlook it. Still it has never been found in Sumatra, Java, Borneo, 
Celebes and the smaller islands of the western portion of the Archipelago, but it 
is the commonest Anopheline in the Moluccas and New Guinea. It is closely 
allied to the Australian N. annulipes and its distribution is in accordance with other 
zoological findings, showing the fauna of the eastern Archipelago to be mainly 
Australian. It is well known that Wallace divided the Archipelago into an Asiatic 
and an Australian region by a line following the Straits of Macassar and of Lombok. 
The Anophelines (and with them other animals ; cf. Weber 1902) do not conform 
to this scheme, as the Anopheline fauna of Celebes, so far as we know at present, 
is purely Asiatic. 


2. Nyssorhynchus schuffneri. 

So far as we yet know, this species is confined to the western provinces of Java 
(Batavia, Bantam) and the neighbouring Lampong districts of South Sumatra, 
which are separated from the former by the Soenda Strait. 


LIST OF THE ANOPHELINES OF THE MALAY ARCHIPELAGO. 91 


3. Myzomyia rossii. 

Christophers (1916) states that M. rossw, Giles, is the common western form, 
whilst M. vaga, Don. (indefinita, Ludl.) occurs throughout Malaya. To this should 
be added that M. rossii does so likewise, as far as the Moluccas ; only in Sumatra 


it has not been found, up till now. 


4. Myzomyia !udlowi and Myzorhynchus sinensis. 

We failed to find both species east of Celebes. But we know that in districts 
where M. ludlowi usually is of common occurrence, it may be absent for months 
and even for a whole year (Citroen 1917). As suitable breeding places were present, 
likewise its common satellite M. rossi, we are not sure that M. ludlowi is really 


absent from the Australian regions of the Archipelago. 


5. Stethomyia aitkenii 
The form with the typical larvae—including all variations described by Stanton 


(1915) and Stickland (1915)—was not found in the Australian region of the Archi- 
pelago. The species was represented there by the varieties insulae-florum and 
papuae, which in their turn were absent from the western regions, except var. 
insulae-florum, which occurs on the south coast of Java (Island of Noesa Kembangan). 

The following table gives a summary of our present knowledge of the distribution 


of the Anophelines in the Malay Archipelago. 


— Ea ae] = a 
3 < © . = ‘a = = 5 = ra ee 2 ee 5 > Pe ies 
zisielnisias| sei Si/./ els esi sls slofgisisisis zlels leas 
VISES Lek | se PS iS i ee | ess is] Si sie s (els) slsell sparsely 
Sir letetsiecls. |e, siealsice sles Stelstele less etsise4ls 
STE S| 5] °S/ 8218/7] 42 Si S818] 8] 8 SST S/S Sle S| Se] ee] § 
< ear ys aiS] siSieit] Jerseys 2 =) 
Sumatra = Sears cere a See te eile aay —— (hy } — 
Java .. vs Sa ee ee se eee tle — 1474] + 
Borneo, incl. findings | 
' ! ily | ' 
of Roper (1914) ../—|—/+}/+/+] + | + /+]/+] + }+/— |4+/-—/+ ne 5-3} 3: 
| 
Celebes a »-(—|—-T-]+]4+] + J + J4+]—] + [4+}/— a8 
Moluccas and New 
Guinea as --}—||—} +4. J] + | + KH — /+} + i ce ee ee ee ee 
Riouw and Linga 
Archipelagos ff —|—|— | +1 — |— sian Himae es banal ep Es ee ee eo 
References. 


CiTROEN (1917). Geneesk. Tijdschr. Ned. Indié, lvii, p. 763. 
CHRISTOPHERS (1916). Ind. Jl. Med. Research, i, p. 454. 

_ Donirz (1902). Zeitschr. f. Hyg. u. Infectionskrankh. xli, p. 15. 
Gries (1900). A Handbook of Gnats or Mosquitoes. London, 374 pp. 
Lup.tow (1904). The Canadian Entomologist, xxxvi, p. 297. 
Mane@kor WinorTo (1918). Geneesk. Tijdschr. Ned. Indié, lvi, p. 462. 
STANTON (1915). Bull. Entomol. Research, vi, p. 159. 


STRICKLAND (1915). Ibid. v, p. 321. 
z (1915). Short key to the Identification of the Larvae of the common 


Anopheline Mosquitoes of the Malay Peninsula. Kuala Lumpur ; 18 pp. 


92 N. H. SWELLENGREBEL AND J. M. H. SWELLENGREBEL DE GRAAF. 


SWELLENGREBEL (1916). De Anophelinen van Nederlandsch Oost-Indié. 
Amsterdam, 182 pp. 

Weser (1902). Die Indo-Australische Archipel und die Geschichte seiner 
Tierwelt. Jena, 46 pp. 


93 


COLLECTIONS RECEIVED. 


The under-mentioned collections were received by the Imperial Bureau of 
Entomology between Ist January and 3lst March, 1920, and the thanks of the 
Managing Committee are tendered to the contributors for their kind assistance :— 

Dr. J. H. Ashworth, F.R.S. :—2 Chironomidae, 16 Culicidae, 6 Tabanus, and 
17 Glossina ; from the Sudan. 

Surg.-Comm. E. L. Atkinson :—4 Tabanus and 3 larvae of Musca domestica. 

Mr. EK. Ballard, Government Entomologist :—46 Culicidae, 478 other Diptera, 
28 Coleoptera, and 47 Rhynchota ; from Madras. 

Capt. P. J. Barraud :—49 Culicidae, 29 other Diptera, 8 Hymenoptera, 377 
Coleoptera, 36 Lepidoptera, and 45 Rhynchota ; from Palestine. 

Mr. G. E. Bodkin, Government Economic Biologist :—6 Tabanidae, 37 other 
Diptera, 350 Hymenoptera, 34 Coleoptera, 100 Isoptera, 1 species of Coccidae, 
51 other Rhynchota, 52 Spiders, 3 Centipedes, and 2 Millipedes; from British 
Guiana. : 

Mr. P. A. Buxton :—14 Trypetidae, 5 Weevils from Cotton, and 1 Bug; from 
Mesopotamia. 

Mr. J. B. Corporaal, Entomologist to the Algemeen Proefstation :—136 Rhynchota ; 
from Sumatra. 

Division of Entomology, Pretoria :—14 Diptera, 189 Hymenoptera, and 7 
Coleoptera ; from South Africa. 

Government Entomologist, Sudan :—3 Diptera, 11 Hymenoptera, 10 Lepidoptera, 
and 5 Rhynchota ; from the Anglo-Kgyptian Sudan. 

Mr. G. F. Hill, Entomologist, Australian Institute of Tropical Medicine :— 
125 Culicidae, 13 Tabanidae, 24 other Diptera, 10 Termites, and 5 species of 
Coccidae ; from Queensland. 

Imperial Department of Agriculture, West Indies :—2 Nests of the Wasp, Polvstes 
annularis and 6 Moths bred from them ; from Barbados. : 

Dr. A. Ingram :—87 Culicidae, 1 Tabanus, 2 Glossina, 15 other Diptera, 1 Moth, 
1 Cockroach, and 2 May-flies ; from the Gold Coast. 

Mr. Nigel K. Jardine :—73 Parasitic Hymenoptera ; from Ceylon. 

Mr. Ll. Lloyd :—Collections of the Tomato Moth, Polia oleracea, L., and the 
Aleurodid, Asterochiton vaporariorum, Westw ; from Hertfordshire. 

Mr. A. Loveridge, Curator, East Africa and Uganda Natural History Society :— 
27 Culicidae, 30 Tabanidae, 20 Glossina, 344 other Diptera, 37 Dipterous pupa cases, 
834 Ants and 37 pupa cases, 6 other Hymenoptera, 1 Caddis-fly, 1 species of Coccidae, 
1,497 other Rhynchota, 2 Orthoptera, and 228 Odonata ; from British East Africa 
and Tanganyika Territory. 

Mrs. W. Smith :—66 Culicidae, 6 Tabanus, 1 Glossina, 50 other Diptera, 21 
Hymenoptera, 23 Coleoptera, 2 Lepidoptera, 2 Caddis-flies, 41 Rhynchota, 6 
Orthoptera, and 1 Tick ; from the Gold Coast. 

Mr. F. V. Theobald :—2 Chalcids (Pteromalus egreqius, Forst.), bred from the 
Brown-tail Moth ; from Romney, Kent, 


94 COLLECTIONS RECEIVED. 


Mr. Robert Veitch :—4 Diptera, 2 Hymenoptera, 3 Coleoptera, and 21 Lepidoptera ; 
from Fiji. 

Mr. Morris N. Watt :—7 Agromyzid Diptera; from New Zealand. 

Wellcome Bureau of Scientific Research :—9 Culicidae, 39 Tabanidae, 1 Hippo- 
boscid, and 23 other Diptera ; from Macedonia and Africa. 

Mr. C. B. Williams :—17 Scolytidae ; from Trinidad. 

Mr. Rodney ©. Wood :—4 Tabanidae, 1 Hippoboscid, 77 other Diptera, 165 
Coleoptera, and 2 Rhynchota ; from Nyasaland. 


a 


IMPERIAL BUREAU OF ENTOMOLOGY. — 
HONORARY COMMITTEE OF MANAGEMENT 


. THE VISCOUNT HARCOURT, Chairman. 
Isgvt.-Coronen A. ALCOOK, OLE., F.RS. | 
Masor KE. E. AUSTEN, D.S.0. Sol ee 
Dr. A. G. BAGSHAWE, O.M.G. | Be Be a sd 
Masor-GENERAL Sir JOHN R. BRADFORD, K.C. M. G., F.R.S. 
Masor-GENERAL Sr DAVID BRUCE, K. O:B., F.R.S. 

Mr. J. CO. F.. FRYER. 
Sir §. F. HARMER, K.B.E., ERS. 
‘Pror. H. MAXWELL LEFROY. 
Hon. E. LUCAS. 
Dr. R. STEWART MACDOUGALL. 
Sm JOHN McFADYEAN, Fe , 
Sir PATRICK MANSON, G.O.M.G., E B.S. 
Siz DANIEL MORRIS, K.0.M.G. 
Pror. R. NEWSTEAD, F.R.S.- 
. Pror. G. H. F- NUTTALL; ¥.RB:8. 
~ Pror. E. B. POU: E.R.S. aS 
Lrzvt,-CoLonet Sir *DAVID PRAIN, OMG. CLE. ERS 
Sir H. J. READ, K.O.M.G., O.B. 
Ta Hon. N. OC. ROTHSOHILD. 
Dr. HUGH SCOTT. 3 | | 
Sim A. E. SHIPLEY, G.B.E., ERS. 5 — 
Mr. R. A. 0. SPERLING, O.M.G. se ae 
Sm STEWART STOCKMAN. | 
Mr. FE. V. THEOBALD. ee ee 
Me, C. WARBURTON. 


&. 


| - . Director. 
Dr. GUY A. K. MARSHALL, OMG. eee aS 
“Bssistant Director. SS Rig ote 
‘Dr. S. As NEAV Bie oo eee es 
Secretary, ee S ‘e 


Mr. Ba: 0:9, PARKINSON. coe - 


~~ \ " . 
a“ 


ie 


95 


OBSERVATIONS ON COTTON THRIPS IN THE GEZIRA, BLUE NILE 
PROVINCE, SUDAN, IN 1918-19. 


By G. H. Corsett, B.Sc. 


From conversations that the writer has had, it would appear that this thrips 
{identified by Mr. R. 8. Bagnall as Heliothrips indicus, Bagn.) has been known on 
the Gezira cotton for about three years, and is popularly called “‘ dry asal”’ in con- 
tradistinction to the work of the “asal fly’ (Aphis sorghi, Theo.) The natives have 
been acquainted with this insect for some time. The reason suggested for the 
enormous damage done by it is that the cotton is too scattered, and it is stated that 
if cotton were cultivated in larger blocks, the thrips might be viewed with equanimity. 

The cotton land at Barakat for this year (1919-20) has been, as far as possible, 
prepared in larger areas, and it will be interesting to compare the result of this 
arrangement with cultivation at Tayiba, where no alteration in grouping has taken 
place. 

Towards the end of October 1918 the thrips attracted serious attention ; by the 
middle of November the cotton looked “ all withered up,” but by the middle of 
December it began to recover. It was not until the beginning of February that work 
on this insect was commenced—most of it being done in the field whilst preparing a 
laboratory to conduct experiments under more immediate control. At that time 
the cotton had practically recovered, in spite of the fact that the thrips was present 
in large numbers. 

In most cases the infested cotton was first noticed on the north side of a “‘ hosa,”’ 
and this implies that the thrips either came from weeds growing on fallow land on 
that side, or was brought froma distance by the prevailing wind, and finding cotton 
and conditions favourable, propagated to such an extent that the immediate study 
of the bionomics and control of this insect was considered necessary. 

The occurrence of this minute insect in enormous numbers caused, during the season 
1918-19, a considerable reduction in the yield of cotton, and it must at present be 
viewed as a great menace to cotton cultivation in the Sudan, though it may prove to 
be quite local in its attack and occur only spasmodically. In any case, at a very 
conservative estimate, an average loss of one and a half kantars of cotton per feddan 
has occurred during the past season on well supervised and cultivated land. 

This is the most important pest with which the cultivators of cotton in the Gezira 
have to contend. On account of its smallness, the rapidity of its spread, and the 
seeming suddenness of its attack in large numbers, the control of the insect is one 
which might present very great difficulties. If the thrips is found on cotton when 
young, it is probably practicable as well as economical to spray ; but on the other 
hand, if 1t comes suddenly when the cotton plants are-about five months old, the 
writer considers the policy of spraying doubtful. 

The injury caused to the cotton plant is due to the larvae and adults feeding on 
both the upper and lower surfaces of the leaves. By piercing the epidermis of the 
leaf and removing the sap, they lower the vitality of the plant. At the points of 
attack spots are formed which, often running together, produce distinct white streaks 

(687) Wt.P2/154. 1,000. 8.20 B.&F.,Ltd. Gp. 11. A 


96 G. H. CORBETT. 


where the chlorophyll has been removed ; and later, if feeding is continued, the lower 
surfaces of the leaves acquire a silvery appearance. As the feeding of this insect is 
prolonged, the leaves become brownish and tough, owing to the efforts of the plant 
to repair the injury. In severe infestations the leaves wither up and fall to the 
ground, preventing the young bolls from developing. 

Leaves frequently present a distorted appearance, curling inwards, and in the 
folds the larvae are often abundant. Larvae have been observed neither on the bolls 
nor in the flowers, but are occasionally seen on the bracts. On plants grown in the 
laboratory, larvae have been noticed feeding on the stem and root just beneath the 
surface of the soil. This undoubtedly was due to the leaves drying up and the thrips. 
migrating for food. In no case was the larval stage completed in such situations. 

Though this insect was first observed as attacking fully-grown cotton trees, it may 
be mentioned that the youngest cotton plants are not immune. For egg-laying, 
preference is shown by the adults for the lower leaves, and then the middle leaves. 
On young succulent leaves at the upper portion of the plant, the thrips are never so 
abundant as they are on the much eaten and tough lower leaves. 


Habits of the Adult. 

The usual mode of progression of the adult is walking, though when disturbed it 
runs, and frequently takes long quick jumps. Flight has been observed in a number 
of cases, but this is not general. Emerging from the pupae in the ground, the adult. 
crawls up the stem to the leaves. In breeding cages it has been noticed to climb up 
the side of the chimney, and when on a level with the leaves to take a leap to the 
plant ; this motion is aided by the wings. 

By far the larger number of the adult thrips feed on and lay their eggs in the tissue 
of the lower surface of the leaf, and confine their attention for the most part to the 
lower and then the middle leaves of the plant. Flowers seem to be immune from 
attack, but larvae have been found on the bracts. Very often the adults are found 
resting and feeding along the veins of the leaves, but, as a rule, no particular part of 
the leaf is preferred. 

Adults enclosed in vials with plugs of cottonwool die rapidly, but under more 
natural conditions they have lived for a maximum of fifteen days, though the average 
works out at five days. The males and females emerge from the pupae about the 
same time, but there are indications that the pupal stage is slightly longer in the males. 
than in the females. 

The female is capable of laying eggs on the second day after emergence. On an 
average, six eggs each day are laid in the tissue of any part of the leaf, but though eggs 
have not been observed, the hatching larvae are more usually found along the veins. 


Habits of the Larva. 

The young larva, on hatching, looks like a starchy-coloured segmented worm, 
with red eyes and with no indications of legs or antennae. Gradually it works itself 
out of the leaf by a slight swaying motion, and standing as it were on its anal 
segment, unfolds antennae and legs. Later, bringing its legs to the leaf surface, it 
pulls itself free. 


OBSERVATIONS ON COTTON THRIPS. 97 


Sometimes it moves only two or three centimetres, but at other times greater 
distances, before commencing to feed. As it imbibes the sap of the plant, the body 
takes on a yellowish hue, and usually by the second day the reddish bands, so very 
characteristic of all stages except the adult, begin to develop. Only one moult has 
been observed—the change from the larva to the prepupa in the soil. 

Like the adult, the larva feeds by piercing and sucking the juices of the plant. It 
prefers the lower surface of the leaf, though it does not by any means confine itself 
there. Movement from one surface of the leaf to the other is common, but from one 
leaf to another is exceptional. Larvae are often found segregated along the veins of 
the leaf, and occasionally in colonies on other parts of the leaf. 

Like many species of thrips, the larva carries a globule of brownish black excreta, 
(on the first day the excreta are colourless), which, after attaining a considerable size 
and dropping on the leaf, dries to a blackish spot. These spots discolour the leaf, and 
their presence is a sure indication that the plant has been, or is being, attacked by 
the thrips. This globule of excreta is held off the surface of the leaf owing to the 
anal segments of the larva being slightly turned upwards. 

The larvae when full grown drop from the leaf, and, crawling into a convenient 
crack in the soil, there change to prepupae. They prefer damp earth, and the 
prepupae are usually found at the junction of the damp and dry soil, which varies 
from about three to six inches down. Immature larvae falling to the ground will die 
if they cannot find food in the nature of weeds or crawl back to the plant, a very 
doubtful possibility. It may be noted that practically all weeds growing amongst 
the cotton support larvae. If the land is kept clean [whilst the cotton crop is 
growing, this will not only prevent the larvae from maturing, but will make the 
adults seek other plants for oviposition. According to experiments, the adults do 
not live more than a day without food. 

The larvae usually like to secrete themselves away from the sun, and are often 
found in large numbers in the curled up parts of the leaf. Larvae in the soil do not 
feed, and, if disturbed, move more quickly than on the leaf. 


Habits of the Prepupa and Pupa. 

The prepupae and pupae are found in the soil, though under unnatural conditions 
they have been observed on four occasions on the leaves. This peculiarity was not 
seen in the field. The larva, before prepupating, does not appear to make any cell, 
but crawls into any convenient place in the soil. The cast skins of the larvae and 
prepupae are found together, indicating that no movement of prepupae and pupae 
takes place. Both stages are capable of movement when disturbed, and are often 
seen in groups, and at other times singly or in pairs. The want of moisture in the 
soil does not have any deleterious effect, but from experiments ‘and observations 
it would appear that if, after the larvae have crawled into the soil, the soil is kept 
moist and not allowed to crack, the adult is unable to emerge. 

The prepupae and pupae possess red bands like those of the larvae ; but whilst in 
the prepupae the antennae are carried forward in front of the head and the wing-lobes 
are partially formed, the antennae of the pupae are bent backwards over the head 
and the wing-lobes are well developed. The darkening of the body of the pupae 
previous to the emergence of the adults is very evident, 

(687) a2 


98 G. H. CORBETT. 


Food-Plants. 

This thrips seems to feed on most plants, cultivated or wild, and below is a list 
of the plants on which the larvae have been okserved. In course of time, no doubt, 
other plants will be found to sustain this insect, and attention may be called to the 
wide range of botanical orders represented in the list. Mr. R. E. Massey, 
Government Botanist, kindly identified most of the wild plants. 


Cultwwated Plants. 

Potatoes, field beans, French beans, butter beans, ground-nuts, wheat, barley, 
peas, turnips, cabbages, cowpeas, bamia, beetroot, sweet potato, lettuce, carrot, 
clover and cotton, both American and Egyptian. 

It has been reported as attacking Lubia (Lubia afin) and Dura, but this the writer 
has not been able to confirm. 

American cotton suffers very much more than the Egyptian, and whilst Eygptian 
cotton recovers remarkably after a severe infestation, in the American variety the 
percentage of recovery is much smaller. 


Wild Plants. 
Malata, the most common plant in the Gezira, generally found with thrips. 
Leucas nubica, Benth. (ungalot) ; very common, and always supporting thrips. 
Phyllanthus mruri, Linn. 
Tephrosia emeroides, Linn. (surep). 
Cucumis melo, Linn. 
Corchorus olitorius Linn. (molokhia). 
Digera arvensis, Forsk. (heyrayrah). 
Solanum incanum, L nn. (gibbein). 
Ipomoea cordofana, Choisy (hantut or tubba). 
Heliotropium europaeum, Linn. 
Heliotropvum supinum, Linn. 
Ocomum basilicum, Linn. (rihan). 
Polygala triflora, Linn. 
Crotalaria sp. 
Abutilon sp. (hambook). 
Abutilon sp. (saphirah). 
Rhyncosia memnonia, Linn. (myoda). 
The local native names for most of the weeds are given in brackets. 


Life-Cycle. 

Observations were made on this insect only from the month of‘November to the 
beginning of August. It is doubtful if there is any aestivating period, and the writer 
is inclined to think that there is no resting stage though, after the rains commenced 
considerable search was needed to find specimens of the insect. The differences in 
the lengths of each stage are remarkable, but under absolute field conditions the 
life-cycle is a short one, and the thrips quickly reproduce in large numbers. 


OBSERVATIONS ON COTTON THRIPS. 99 


As the result of a number of observations and experiments it was found that the 
length of the egg stage varied from 4 to 15 days, with an average of 8°2. In February 
the minimum and maximum were, respectively, 9 and 15 days; in March, 6 and 11; 
and in April, 4 and 9. 

The period during which the larva remained on the plant varied from 3 to 6 days, 
with an average of 4:2. From 4 to 14 days elapsed between the entrance of larvae 
into the soil and the emergence of the adults, the greatest number of emergences 
taking place on the sixth day. 

From this it will be seen that the developmental cycle may be as short as 11 days, 
or may take as long as 35 days; but the writer is of opinion that 18 days may be 
regarded as the usual time, and the adults do not begin egg-laying until two days 
after their emergence from the soil. 


Control Measures. 

In discussing the control of this thrips, it should be borne in mind that its be- 
haviour after the old cotton crop is removed till the new one is well advanced is not 
yet precisely known. Larvae were difficult to find on wild plants at the beginning 
of August, but until the rains commenced they were numerous. 

Control may be considered under three heads, namely, suggestions, cultural 
methods, and insecticide applications. 

1. Suggestions. 

(a). If this insect spends the “ dead season” on vegetation, all plants of every 
description should be removed ; but if, on the other hand, it rests in the soil, the 
ploughing of the old cotton land should take place as soon as convenient so as to 
expose the aestivating stage. 

(6). Larvae and adults are very numerous on the last standing cotton, and the 
writer suggests that, since there is so little to be gained and so much to lose, the 
watering of the cotton should cease slightly earlier and the cotton should be cut out 
and burnt sooner; by this means the number of thrips that could attack the suc- 
ceeding cotton crop would be greatly diminished. 

(c). Observations tend to show that heavy waterings have a detrimental effect on 
the numbers of thrips emerging from the soil. The probable reason for this is to be 
found in the fact that the soil does not crack so quickly, and the adults, unable to 
_ escape from the ground, succumb. The effect of irrigation on this thrips is a line on 
which further investigations are required. It is suggested that one or two heavy 
waterings about the beginning of November might have a very beneficial effect. 

(d). Owing to the wide range of food-plants, experiments with trap crops should 
be tried. 

2. Cultural Methods. 

The cultivators of cotton in the Gezira appreciate the extent of the damage this 
insect can do, and the destruction of all weeds should be the first preventive 
measure to be adopted. Practically every plant in the vicinity of cotton has been 
found to harbour the larvae, and all vegetation growing amongst the cotton or in its 
vicinity should therefore be eradicated. This not only applies to the growing season, 
but is equally applicable to land which will be under cotton the following year, and 
to the banks of all canals and gadwels. 


> 


100 G. H. CORBETT. 


It should always be borne in mind that clean cultivation not only prevents, in» 
many cases, the completion of the life-history of this insect, but also yields other 
advantages to the crops. 


3. Insecticide Applications. 

Four insecticides were experimented with, namely, pyridine sulphate, quinoline 
sulphate, nicotine sulphate, and soap. 

Pyridine sulphate was found unsuccessful. 

Quinoline sulphate possesses insecticidal properties, but the fact that it had to be 
used in strong solution, and therefore in large quantities, rendered it unsuitable. 

Nicotine sulphate was tried with soap (“‘ Sunlight ’’) in varying strengths, and the 
best solution found was one part of 32 per cent. nicotine sulphate to 1,200 parts of the 
soap solution, which was obtained by dissolving 1 lb. of the soap in 21 gallons of water. 

Soap (“Sunlight’’) spray, consisting of one pound of soap to ten gallons of water, 
will kill a large percentage of thrips, both larvae and adults. Soap alone is only 
recommended if nicotine sulphate is unobtainable. 

It should be stated that spraying is considered impracticable, as well as unecono- 
mical, when the cotton is four or five months old; but if the thrips should appear 
when the plant is small, the nicotine sulphate spray with soap will be found very 
useful. Spraying should be done in the late afternoon and evening, and at least 
twice, so as to kill the larvae which have hatched from eggs in the leaves since the 
first spraying. 


101 


SOME FURTHER NOTES ON THE TSETSE-FLIES OF NYASALAND. 
By W. A. Lamporn, M.R.C.S., L.R.C.P. | 


Glossina morsitans. 

A fortunate stay of a few days in July 1919 at Fort Johnston, Lake Nyasa, 
while waiting instructions, enabled the writer to proceed to the village of Mzeze 
near the Livingstonia Peninsula, where in 1914-15 some work was carried out on the 
various insects parasitic on the puparia of this fly. 

The success at that time of the various parasites, especially Mutilla glossinae, 
Turner, as judged by the numbers of their cocoons found in old puparia of the fly and 
bred from recent ones, had been phenomenal, and so the chance, though one lasting 
only a few hours, of ascertaining the progress made by these insects in the course of 
the three subsequent years was a very welcome one. 

The early rainy season was quite similar in character to that of 1915, when long 
series of the flies had readily been captured in the district, on passing through which 
one had been constantly harassed by a buzzing swarm. Far different was one’s 
present experience, for though the journey was made on a cycle, which attracts more 
flies than a pedestrian, none at all appeared, and it was only on the following day 
that a single specimen was taken in the course of a hunt for puparia. Local natives 
agreed as to the diminution in numbers of the flies thereabouts without there being 
any falling off of game, and added that the pests had now become more numerous to 
the north, a statement substantiated by personal experience a few weeks later on. 

A gang of six boys, who had on a previous occasion worked as puparia collectors 
and were therefore experts, were got together to assist again in collecting puparia, 
and the results were confirmatory ; for though two days were devoted to the search, 
only 107 old puparia and 15 new ones were forthcoming, mostly from the artificial 
breeding places designed in 1915. Examination of the old puparia showed that nine 
definitely had been destroyed by Mutillid parasites, but it was not possible 
to determine with certainty the full extent of the parasitism either by this species 
or others, owing to the age and broken condition of many of the puparia. Of the 
fifteen new puparia three only afforded flies, a male on 23rd August, a female 
on lst September, and another female on the 4th. Five yielded Mutilla glossinae, 
two males on 17th September, one male on 18th, and one of each sex as long after 
as 29th. The remaining puparia all died, possibly as a result of the vicissitudes of 
temperature to which they were exposed, for they were taken on a month’s foot 
journey in the uplands of Angoniland, at an average elevation of from 3,000 to 
4,000 feet, hot days being generally succeeded by chilly nights. 

Whereas in 1915 at Mzeze parasites would be readily obtained, at Domira Bay 
none at all were either captured or bred out, this being about fifty miles to the north 
and separated from the Mzeze area by the south-west arm of Lake Nyasa. No 
opportunity of visiting the area at Domira Bay occurred, but in September 1919 
a visit was made to another section of it, at Kotakota, about forty miles north of 
Domira Bay, where between the 6th and 10th of the month hundreds of old 
G. morsitans puparia, and sixty-five new ones were obtained. None of these old 
pupa-cases showed the Mutillid cocoons, nor did any of these parasites emerge from 


102 WwW. A. LAMBORN. 


the new puparia, of which twenty-six yielded flies (twelve males and fourteen 
females) during late September and the first week of October, the remainder dying. 

The morsitans area in the neighbourhood of the Livingstonia Peninsula and 
running inland for miles, so as probably to be continuous with the great area running 
down the west bank of the bed of the former Shire River, had undoubtedly been 
infested with these flies for many years, various natives, well on in years, stating 
that they remember them even when they were children ; whereas the area running 
from Domira Bay up north to Kotakota, and beyond it, and south towards Dedza, 
in which the fly is far more abundant, has undoubtedly been more recently occupied 
by them; for many of the younger Government officials remember when large 
herds of cattle grazed on its wide dambos, now entirely destitute of all domestic 
animals, though there are large settlements of natives. The evidence would seem 
to suggest that the abundance of the fly thereabouts may be due to the failure of 
the natural parasites to keep equal pace with the fly in the migration which has 
occurred. 


Glossina brevipalpis. 

This species was found by the writer in the same area with morsitans in 
the course of a journey through a stretch of twenty miles of country between 
the mouth of the Bua River, twelve miles north of Kotakota, and that of the 
Duangwa, in every ravine and hollow where large shade trees were associated 
with heavy undergrowth. As showing how readily the presence of this fly may be 
overlooked it may be remarked that on the outward journey not one was seen, 
and that it was only on the return that the capture of two males in the early morning 
suggested a search. Many were then taken at dusk, and the numbers of the puparia 
obtained will show how numerous the flies really must have been. 

On the 12th of the month a systematic search for their breeding places was com- 
menced. A little scratching with a bit of stick in the loose sandy soil, much lightened 
by humus, among the herbaceous vegetation beneath a huge shade tree soon brought 
to light twenty pupa-cases ; more were found during the pitching of the tent, anda _ 
gang of raw natives who had been attentive to the mode of search were then put 
to the work. Within four hours the most industrious man had got together 390 
puparia (old and new), two others bringing in 267 and 257 respectively, and between 
the 12th and 15th a total of no less than 13,838 old puparia, with 248 new ones, 
were found by them. At the same time in these breeding places only 23 puparia 
(17 old and 6 new) of G. morsitans were found, showing how different are the 
breeding places selected, for morsitans was present in swarms. 

Twelve of the old puparia of G. brevipalpis were whole except for a small puncture 
_which had probably given exit to parasites, though neither on this occasion, nor 
when the puparia were previously studied, were any of these obtained. The 
perforations were of two kinds, one about the size of the head of a small pin and 
situated usually midway between the two poles, and the other a much larger breach 
less regularly rounded and more commonly towards one or other pole. Twenty- 
eight only of the new puparia yielded flies (13 males and 15 females), and all the rest 
perished. All these flies came out an hour or two before dusk, except five, all males 
which emerged, four at about 10 a.m., one at 2 p.m. 


NOTES ON TSETSE-FLIES OF NYASALAND. 103 


On two successive nights camp was pitched in localities where these pupae were 
being obtained with a view to ascertaining whether brevipalpis is nocturnal as well 
as crepuscular in its habits. Though 32, all males, were taken at dusk, between 
6 and 6.15 p.m., all within 50 yards of the tent, two only were taken inside it. 
None were taken after dusk, though the nights were bright and starlit, nor were they 
attracted to the camp lamps. At about 8 p.m. a search was made in the bush by 
the light of a bullseye lantern, but again the flies were neither seen nor heard ; and 
though the party of 23 natives slept out in the open, few having blankets, none 
complained of having been troubled by the fly. 

The breeding grounds of this fly were further studied in October, some two 
hundred miles north in the vicinity of the Lufira River, North Nyasa, close to the 
Lake, a locality long notorious for the fly, the places selected by it having the same 
character as all the breeding places previously examined. In this neighbourhood 
13 new puparia and 9,094 old ones were collected between the 7th and 18th of the 
month, only 14 of which showed perforations suggesting the exit of parasites. 

In the course of the journey through the fly-infested villages north of Kotakota 
the method was witnessed by which the natives of that part, while sitting down, 
catch, with the least effort to themselves, flies settled within reach on their bodies. 
The blade of an old sheath-knife or an old spear blade is placed almost flat on the 
skin and is then advanced, with the edge slightly raised, by no means slowly or even 
very carefully, towards the fly, which, though very alert in regard to menace from 
above, evinces no apprehension of that below it. When the edge is well over its 
feet the native presses them on to his skin with it, and the fly, so trapped, is then 
taken in the fingers and subjected to treatment such as is doubtless considered 
justifiable by reason of the annoyance it has caused. © 


Suggestions in regard to Tsetse Control. 

One of the most pressing entomological problems at the present day in the British 
Empire is that of the control of the various species of tsetse-flies. As shown by a 
number of workers, the parasites destructive to their puparia are not few, at all 
events in the case of G. morsitans, and in some areas these parasites are known to 
exert a considerable influence in reducing the numbers of the fly. But, generally 
speaking, no very material reduction, from the point of view of man, would seem 
to be effected by these agencies; though at this early stage of research in regard 
to the fly it may perhaps be premature to make any unqualified statement as to their 
control value. Instances have been recorded where the tsetse-fly in a district has 
unaccountably diminished without there being any marked diminution in the numbers 
of the game animals therein. Major E. E. Austen in his ‘‘ Handbook of the Tsetse- 
flies ’’ (1911, p. 65) mentions the disappearance of G'. morsitans from the Victoria 
Falls, where at one time it abounded. The late Captain F. C. Selous, in his book 
“ African Nature Notes,” speaks of the disappearance of the flies at the same time 
with buffalos from the valley of the central Limpopo and its tributaries, where 
other game—kudu, zebra, wildebeeste, hartebeeste, impala and bushbuck— 
continued to exist in considerable numbers, and he suggests that the flies died out 
because they were unable to maintain themselves on game other than buffaloes. 
More recent study of the flies has shown that this is not the case, and the writer 


104 W. A. LAMBORN. 


suggests that a more reasonable explanation is to be found in the local extermination 
of the flies by their parasites, a process, as he believes, now steadily proceeding in 
the area near Mzeze, Nyasaland. The question therefore naturally arises—how 
far is it possible to increase their influence either by the introduction of new parasites 
or by breeding on a larger scale those already known to science ? 

The genus Glossina being now limited to the Ethiopian region, it is doubtful how 
far the parasites of other Diptera, brought in from other lands, would seek out and 
destroy its puparia. It is to be apprehended that those obtained from the puparia 
of other Muscids, breeding under more or less similar conditions, might doso. A more 
hopeful line of action could possibly be found in the search for other species naturally 
parasitic on Glossina but having a different geographical distribution. For instance 
a Bombylid fly, Villa lloydi, Austen, parasitic on G. morsitans, has so far only been 
discovered in Rhodesia, and a second species of parasitic Mutillid, MW. benefactriz, 
Turner, in Nyasaland only. ‘The parasites of the fly on the West Coast are probably 
different from those in the East and South, the insect fauna, generally speaking, 
being largely distinct. The various species of tsetse-flies may each have different 
parasites yet to be discovered, which might be interchangeable. 

It is probable, and some of the evidence on this point is very definite, that in the 
case of rapid extension of a fly area, parasites less endowed than their hosts with the 
power of ranging far and wide—strength on the wing having doubtless played a 
considerable part in the present-day success of the tsetse as an insect—have 
lagged more or less behind in the foci first inhabited by the flies. And this is parti- 
cularly likely to be the case with Mutillids, the females of which are wingless. 
One of the present suggestions in connection with fly control is to endeavour to 
enhance the value of the parasites already known to science, the Chalcids in parti- 
cular. They could unquestionably be bred readily enough in the laboratory on the 
puparia of their natural host, though owing to the difficulty im obtaining these 
in any numbers, the laboratory output would necessarily be very limited. It has 
occurred to the writer that it might be possible to employ as alternative hosts the 
puparia of some of the common Muscids, then liberating in great numbers the little 
insects, each with its dominant aim in life to seek out and destroy the greatest of 
the insect scourges of man and beast in the African Tropics. 

The general upset of work as a consequence of war conditions made any attempt 
to put the idea to a practical test impossible, but an opportunity was made to carry 
out, during 1918 in Kast Africa and during the last six months of 1919 in Nyasaland, 
research work in regard to the question of alternative hosts for Chalcids normally 
parasitic on certain Muscids. The results (which will be written up later) showed 
some promise, though this work has again, unfortunately, been brought to a prema- 
ture close. 


105 


THE EARLY STAGES OF WEST AFRICAN MOSQUITOS. V.—CULEX 
DECENS, THEO. AND CULEX INVIDIOSUS, THEO. 


By J. W. S. Maorie and A. InerRam, 
West African Medical Service. 


Both Culex decens and Culex wnvidiosus are widely distributed in British West 
Africa. In the Gold Coast both have been taken in all the three divisions into which 
the country is divided, namely, the Colony proper, Ashanti and the Northern 
Territories ; the records at Accra showing the following distribution :— 


Culex decens. 

Colony—Accra, Cape Coast Castle. 

Ashanti—Kintampo, Kumasi, Sunyani. 

Northern Territories—Batiasan, Bawku, Binduri, Bole, Daweni, Dogan Kade, 
Jefisi, Kalleo, Kpalgu, Lorha, Mayoro, Nandaw, Nandawli, Nasia, Navarro, 
Pinna, Salaga, Sambolugu, Savelugu, Tamale, Tanina, Tiana, Tishi, Ulu, 
Wa, Yaga, Zuaragu. 

Culex invidiosus. 
Colony—Accra, Akuse, Bibianaha, Koforidua, Nsawam, Sekondi. 
Ashanti—Akrokerri, Bjere, Kumasi, Obuasi, Odumase, Sunyani. 


Northern Territories—Bawku, Bole, Gambaga, Kugri, Lorha, Maliki’s Zongo, 
Navarro, Tumu, Turu, Wa, Zuaragu. 


So far as they go these records do not point to any cleavage in the areas of distri- 
bution of the two mosquitos, both species having been collected in a number of the 
same places. 

The majority of the records for the Northern Territories were made by one of us 
(A.I.) during a tour undertaken in 1918, the observations made at this time suggested 
that C. decens might be a domestic variety of C. invidiosus. Edwards (Bull. Ent. 
Res. ui, p. 381), however, considers the two to be distinct. In the adult, he says, 
““ C. decens can be distinguished by the reddish thorax (that of C. invidiosus being 
brownish), and (in the male) the banded abdomen. In the female the abdominal 
banding is not constant ; the bands in C. decens are always narrow and may some- 
times be interrupted.” But he failed to detect any differences in the genitalia of 
the males (an observation which he repeated later, Bull. Ent. Res., v, p. 70), and was 
unable to separate the larvae, considerations which led him to admit that it was 
** quite possible that the two are really only forms of one species.” 

We have compared a number of larvae and pupae of these mosquitos in an 
endeavour to find some means of distinguishing them. We may say at once that we 


106 J. W. S. MACFIE AND A. INGRAM. 


were unable to detect in the larvae any constant difference of specific importance, 
and as this observation is in accord with that of Edwards further details will not 
be given. 

In a previous paper (Bull. Ent. Res. vin, p. 85) we have described the pupa of 
C. nvidiosus. The pupa of C. decens (= C. mgrocostalis, Theo., and C. lividocostalis, 
Graham) has been briefly described by Wesché (Bull. Ent. Res. 1, pp. 40 and 45), 
but the details given are meagre. As it was necessary on this account to re-describe 
the pupa of C. decens, and as it seemed advisable to give at the same time further 
details regarding C. invidiosus because of the very close resemblance of the two 
pupae, we decided to compare seta by seta five specimens of each. The result of 
this examination will be recorded here in detail. 

The pupae were found to be similar both in their general features and as regards 
the arrangement of their cephalo-thoracic and abdominal setae. In describing 
the setae the same method has been adopted as that employed by one of us in des- 
cribing the pupa of Stegomyza fasciata (Bull. Ent. Res. x, pp. 161-172). 


General Features. 

In Table I the general features of the pupae are tabulated. The figures given are 
the averages of the measurements made in each case. The specimens of C. invidiosus 
examined were on the whole slightly smaller than those of C. decens. It will be 
observed that the general features of the two pupae were similar. 


TABLE I. 
General Features of the Pupae of C. decens and C. invidiosus. 


OC. decens C. invidiosus. 


Length of pupa, extended .. | 4-44 mm. 4-44 mm. 
Respiratory trumpet, 
total length 640u 580u 
length of the open ‘portion 
(pinna) 170u 160u 

ratio of the ‘length of the 
closed portion (meatus) 

to the total length ee a alee 1:14 

Length of the tuft (A) at the 
posterior angle of the 

seventh abdominal seg- 


ment ee ae at aOR 1904 
Length of the tuft (A) at the pos- 

terior angle of the eighth 

abdominal segment .. | 210u 190u 
Paddle, length .. mn .--| 750n 740u 

breadth .. 530u 530u 

ratio of length to breadth | 1:4: 1 L431 


Cephalo-thoracic Setae. 

The setae on the cephalo-thorax of these pupae are alike, and are similarly situated 
to those of Stegomyia fasciata (loc. cit.) ; they do not require special description. 
Details regarding the subdivisions of the cephalo-thoracic setae are furnished in the 
following comparative table (Table II). 


THE EARLY STAGES OF WEST AFRICAN MOSQUITOS. 107 


Fig. 1. Diagram showing the arrangement of the setae on 
the cephalothorax of the pupa of Culex decens or CO. invidiosus. 


TaBLeE II. 
A Comparison of the Cephalo-thoracic Setae of the Pupae of C. decens and C. invidiosus. 


| 


Seta. OC. decens. C. invidiosus. 


Post-ocular, | 
Superior ; rather long, not 
especially strong ; 3 | 3-4, usually 3 
Median; rather Tong, not 
especially strong 4-5, usually 4 4—5, usually 4 
Inferior ; rather long, not | 
especially strong .. | 3-4 3 
Antero-thoracic, 
Lower anterior; moderate 
length, constituent hairs 


rather delicate .. 4-5 4-7 
Upper anterior; rather 
small and delicate on tee 3 
Lower posterior ; long, but 
not very strong .. Ry te 2-3, usually 2 


Upper posterior; rather 
small and delicate, 
situated above the 


lower posterior seta .. | 1-4 2-5 
Dorsal; moderate length and 

Strength . 2-4, usually 4 3-5 
Supra-alar ; moderate length and 

strength . .s spine 2-3, usually 2 


Postero-thoracic, 
Internal; well-developed 
tuft of moderate length 


and strength .. 5-8 5-8 
Median; rather long and 
strong . 2-3, usually 2 y 


External ; tuft of moderate 
length ‘and strength, or 
rather long tg 3- 


Cr 
he 
Or 


| 
| 


108 J. W. 8S. MACFIE AND A. INGRAM. 


Dorsal Abdominal Setae. 

The setae on the dorsal and lateral aspects of the abdomen may be described 
together. Details as to the subdivisions of these setae will be found in the tabular 
statement which follows; the facts given here are either of a general nature, or 
such as may be required to make the table comprehensible. 

The following setae, which form series, are recognisable on each side of a typical 
segment of either pupa :— 

1.—The lateral seta (A), situated a little above and internal to the posterior angle. 

2.—The seta belonging to the sub-median row (C), on the dorsum, situated near the 
posterior margin of the segment about half way between the posterior angle and the 
middle line of the abdomen. This is the row of setae which on a previous occasion 
(Bull. Ent. Res., x, p. 59) was referred to as the wnner lateral row. 

3.—A very small seta (C’), situated near the posterior margin of the segment and a 
little internal to C. . 


Fig. 2. Diagram showing the arrangement of the setae on the dorsal and ventral aspects 
of the pupa of Culex decens or C. invidiosus; dorsal setae shown on the upper, ventral 
setae on the lower half. 


4.—The seta belonging to the sub-lateral row (B) on the dorsum, situated near the 
posterior margin of the segment about half way between the posterior angle and 
C. This is the row of setae which on a previous occasion (Bull. Ent. Res., x, p. 59) 
was referred to as the outer lateral row. 

5.—Two setae (B’ and B”) situated a little anterior to the posterior margin of the 
segment ; B’ outermost and always more or less external to B, B” internal to B’ and 
on some segments external, on others internal to B. 3B” is sometimes associated 
more closely with C than with B, for example on the sixth segment. . 

6.—A very minute seta, the anterior dorso-central seta (D), near the anterior margin 
of the segment in line with the setae of the sub-median row. 

These seven setae are easily recognised on the third to the sixth segments; the 
setae on the other segments do not conform completely to this plan and require a 
few words of explanation. 

Segment I. The setae on this segment are arranged in a similar manner to the 
corresponding setae of Stegomyia fasciata, and the same names are used in describing 
them, namely, Dendritic tuft for the large branched seta in the middle of the segment 
near the median line, and, for the setae along the proximal border, from within 
outwards, Antero-internal, Antero-external, Medio-internal, Medio-external, Postero- 
internal, Postero-external and Lateral. 


THE EARLY STAGES OF WEST AFRICAN MOSQUITOS. 109 


Segment II. Nine setae may be seen on this segment in a dorsal view, of which 
C and D are as defined above. Internal to C, near the posterior margin of the seg- 
ment, is a small dendritic tuft (D. T.). The setae external to C are as follows: a 
small single seta (B) a little external and anterior to C, a tuft of moderate size (B”) a 
little external and anterior to B, and a small tuft (B’) a little external and posterior 
to B”. Laterally, or ventro-laterally, there are three setae: the most anterior (A’) a 
very long, stout seta ; rather posterior to A’ a seta (A) similar to the Lateral setae on 
the more posterior segments ; and posterior to A a very small seta (S) similar in 
appearance to the Postero-lateral setae on the ventral aspect of the abdomen. 

Segment VII. A, B, B’, B’, C, and D as defined above. There is no small seta 
internal to C, but a little external to it there is asmall seta similar to C’ on the other 
segments ; this seta is referred to in the Table as C’. In addition there is a small 
tuft (A’) close to and a little posterior to A. 


Segment VIII. A and D as defined above. 
(P) situated close to the external margin of the root of the paddle. 


In addition there is a delicate seta 


At the distal end 


of the midrib of the paddle there are two very small setae (P’ and P”) ; usually single, 


but sometimes forked at their ends or even divided more deeply. 


TaB.eE IIT. 
A Comparison of the Dorsal Abdominal Setae of the Pupae of C. decens and C. invidiosus. 
a Seta. O. decens. O. invidiosus. 


I. |Dendritic tuft ; 


large and well- 


developed, strong primary branches | 8-12 8-11 
Antero-internal ; short, rather stout | 1, end occasionally 1 
forked. 
Antero-external; rather long and 
strong be ls na .. | 2-3, usually 2 2 
Medio-internal ; small ne .. | tora amalltatt* 1, or a small tuft 
Medio-external; small, but longer | Tuft of about 2-5 Sunilar tuft 
than the Medio-internal oii hairs. 
Postero-internal; very long, strong | 1-2, usually 1 1 
Postero-external; rather long and 
slender - ee dy? 2 1-3 
Lateral; small, delicate 1, end sometimes | 1, end sometimes 
divided. divided. 
II. |A_ ; rather long and slender. . Z 1-3 
A’ ; very long, strong 1 1-2 
S ; very small and feebly chitinised | 1 1 
B_; small, rather stout > e | 1 
B’ ; small tuft 7 Bs .. | Smalltuftof2—5hairs) Similar small tuft 
B’ ; small tuft, but considerably 
larger than B’, situated between 
B and B’ and anterior to them | Tuft of 4-6 hairs Tuft of 4-6 hairs 
C ; moderate length, reaching nearly 
half way across the third ae 
ment. oa 2-3, usually 2 2 
D.T. ; small dendritic ‘tuft Small dendritic tuft | Small dendritic tuft 
D ; minute ; 1 1 
III. |A_ ; rather long and slender 2-3 2-4 
B_ ; rather long and strong .. ra 2 
B’ ; small tuft between A and B” .. | Smalltuft, 1-7 hairs) Small tuft, 1—4 hairs 
B’” ; 


tuft, considerably longer than B’, 
situated between B and B’ .. 


Tuft, 3-7 hairs 


Similar tuft 


*In the case of small tufts, the number of constituent hairs, when given, is only approxi- 
mate as it was often impossible to determine with accuracy the degree of sub-division. 


110 


J. W. S. MACFIE AND A. INGRAM. 


sch Seta. C. decens. C. invidiosus. 
C ; well-developed tuft, reaching | 
about half-way across the fourth 
segment a: Be 6-9 6-12 
C’ ; small AG ee 1 
D ; minute i « Wal 1 
Iv. |A_; rather long and stoner 2-4 3-4 
B_ ; long, strong, reaching almost 
across the fifth segment 3-4 4—6 
B’ ; small tuft, external to B Small tuft, 1-3 hairs | Small tuft, 1-3 hairs 
B’” ; small tuft, anterior to B : Small tuft, 4-6 hairs | Small tuft, 4~7 hairs 
C ; well- developed tuft reaching fur- 
ther than half-way across the 
fifth segment 4-8 6-10 
C’ ; small 1 1 
D ; minute I 1 1 
Vv. |A_ ; rather long and slender . 2 3-4 3-4 
B_ ; very long and strong, reaching 
beyond the posterior margin of 
the sixth segment 5 2 2 
B’ ; rather small tuft, external to By Small tuft, 2-5 hairs | Small tuft, 2-6 hairs 
B’ ; moderate length, external and 
anterior toC .. 1—2, usually 2 2-3, usually 2 
C 3; well-developed tuft, reaching fur- 
ther than half-way across the 
sixth segment 4-8 6-7 
iC”. svsmall 1 1 
D ; minute 1 1 
VI. |A_ ; rather long and alontes 3-4 3-5 
B_ ; very long and strong, reaching 
beyond the posterior margin of 
the seventh segment sia Ae 2 
B’ ; small tuft, external to B 2-4 2-4 
B” ; moderate length, situated exter- 
nal and anterior to C 2 2 
C ; well-developed tuft, reaching 
about half-way across the 
seventh segment : 4-7 4-8 
C’ ; small i 1 
D ; minute : = : i 1 
VII. A ; constituent hairs strong, usually 
sub-plumose and often branched, 
sometimes simple : 3-5 , 2-4 
A’ ; small tuft ie Small tuft Small tuft 
B_; moderate length and strength . 2 1-3 
B’ ; moderate length, external to B. 2 2 
B’” ; moderate length, internal to Bo. ees 2-5 
C ; moderate length, reaching about 
half-way across the eo Seg: 
ment... 2-5 4-7 
C’ ; small, external to C 1 1 
D ; minute : 1 1 
VIII. |A ; well-developed tutt, ame anes 
hairs strong, u usually branched 
and sub- aaa t 6-7 5-9 
D «-minute’— ; 1 1 
P ; moderate length, ‘slender 2 2 
P’ and P”; very small Two small setae Two small setae 


THE EARLY STAGES OF WEST AFRICAN MOSQUITOS. Lt 


Ventral Abdominal Setae. 
The setae present on each segment, as well as the characters of the setae, are shown 
in the following tabular statement (Table IV). 
TABLE IV. 
A Comparison of the Ventral Abdominal Setae of the Pupae of C. decens and C. invidiosus. 


Seg- Seta. C. decens. CO. invidiosus. 
ment. 
III. | D ; very small, feebly chitinised ../ 1 1 
E_ ; small tuft, longer than E’ Small tuft, 4-5 hairs | Small tuft, 2—5 hairs 
E’ ; small tuft Small tuft, 2-5 hairs | Small tuft, 4-5 hairs 
C ; moderate length 2 ; 2 
B’ ; small 1 1 
A; minute 1 1 
IV. | D ; very small, feebly chitinised 1 1 
fos sroall tuit. Ms 1, or small tuft, 2-5 | Small tuft, 2-4 hairs 
hairs. 
E’ ; small tuft, longer than E 1, or small tuft, 2-6 | 1, orsmall tuft, 2-4 
hairs. hairs. 
C ; moderate length 
B’ ; small t 1 | 
A ; minute 1 1 
V. | D ; very small, feebly chitinised 1 1 
E ; small tuft ah it 1, or small tuft, 2-4 | Small tuft, 2-4 hairs 
hairs. 
E’ ; small tuft, longer than E Small tuft, 2-5 hairs | Small tuft, 4~7 hairs 
C ; rather long, reaching fully half 
way across the sixth segment ..| 1 1 
B’ ; small a 1 1, end sometimes 
forked. 
A ; minute 1 1 
VI. | D ; very small, feebly chitinised 1 1 
E ; small .¢ is 1, or small tuft, 2-4 | 1, or small tuft, 2-3 
hairs. hairs. 
C ; rather long, reaching fully half 
way across the seventh segment} 1 1 
B_; rather long, shorter than C 1 1 
B’ ; small t 1-2, usually 1 1-2, usually 1 
A : minute .. 1 1 
VII. | E ; small tuft Small tuft, 2-4 hairs | Small tuft, 2-4 hairs 
C ; rather long, reaching fully half 
way across the eighth oo ] 1 
B_; moderate length 1 1 
B’ ; small, end sometimes forked or 
sub- divided, a little external 
and anterior to B 1-2 1—4, usually 1 
A ; minute 1 1 
Vath, | A’ ;. minute 1 1 


The setae on the ventral aspect of the abdomen are few, and mostly inconspicuous. 
The following setae, which form series, may be recognised on each side of the abdomen 
of either pupa on two or more of the segments :— 

1.—A small seta, the medio-lateral seta (E), a little internal to the lateral border and 
rather posterior (junction of the posterior and middle thirds) to the middle of the 
segment. 


(687) 


112 J. W. 8S. MACFIE AND A. INGRAM. 


2.—A very small seta, the postero-lateral seta (D), in a lateral position just above 
the posterior angle. 

3.—A small seta (E’) a little posterior and external to E. 

4.—The seta belonging to the inner ventral row (B). 

5.—The seta belonging to the outer ventral row (C). 

6.—A small seta (B’) near the posterior margin of the segment a little internal to C. 

7.—A very minute seta, the anterior ventro-central seta (A), at the anterior margin 
of the segment and very close to the middle line of the abdomen. 

Two setae require special mention. On the eighth segment there is a minute seta 
(A’) near the anterior margin of the segment a little internal to the level of B on the 
seventh segment. This seta resembles A, but is placed more externally and pos- 
teriorly than A on the more anterior segments. On the seventh segment the small 
seta internal to C is situated close by and a little anterior to B; we have therefore 
marked it B’, and have retained this symbol for the similar setae on the more anterior 
segments which, however, are situated nearer the posterior margin and more exter- 
nally. 

It is clear from the descriptions which have just been given that the setae of the 
pupae of C. decens and C. invidiosus are similar in character and situation, and that 
although they are somewhat variable as regards their sub-divisions the degrees of 
variation of the two overlap. In fact, they provide no feature of differential im- 
portance. 

The question then arises, are C. decens and C. wnvidiosus separate species or are they 
varieties of a single mosquito. There are indeed certain differences in the adults, 
but the genitalia of the males are identical according to Edwards, the larvae cannot 
be separated, and the same remark applies to the pupae. Under these circumstances 
we think there can be little doubt that they should be regarded as varieties and not 
as distinct species ; we propose to retain for the species the name C. decens. 


113 


SOME NOTES AND REMARKS ON THE BIONOMICS OF GZOSSINA 
MORSITANS. 
_By Rupert W. Jack, F.ES., 
Agricultural Entomologist, S. Rhodesia. 

It is with considerable diffidence that the writer has prepared the following paper, 
not only on account of several previous contributions to the subject, but also, and 
chiefly, because no opportunity has been available for more than comparatively 
brief visits to the fly areas in Southern Rhodesia, and the observations to be recorded 
are therefore of a somewhat meagre nature compared with work that has been carried 
out of recent years in other parts of Africa. The tsetse-fly problem is, however, of 
such paramount importance to Africa as a whole, and the continued extension of the 
fly belts such a grave hindrance to the full development of Southern Rhodesia, that 
it is felt that too much time and thought cannot be devoted to the subject, and that 
every contribution whether of actual observations or of theoretical deductions, or of 
both, must have a certain value, if only in provoking discussion and criticism. In 
the following pages the important question of whether or not Glossina morsitans is 
vitally dependent upon the larger mammals is not directly dealt with. The strong 
trend of scientific opinion in the direction of an affirmative conclusion must be 
admitted at the present day by the most vigorous opponent of the theory. There is 
hardly an investigator of any standing who has not contributed some observations 
of weight, indicating that the fly is mainly dependent upon the larger mammals for 
its food supply. For the purposes of the present discussion this dependence has been 
to a large extent assumed with a view to an attempt to explain certain phenomena 
upon this basis. The discussion is, however, of a very disconnected nature and aims 
merely at touching upon a few points in connection with the bionomics of the fly that 
need clearing up, and if the writer’s views provoke criticism, or even direct con- 
tradiction, supported by actual observations, the paper will have served its purpose, 
as our knowledge of the subject cannot fail to benefit thereby. 

It will be found that there is a very substantial agreement between observations 
in this territory and those recorded further north, particularly in Northern Rhodesia, 
but there are several important points bearing on the fly problem concerning which 
a considerable difference of opinion appears to exist, and others that have not 
apparently been touched upon as yet. 

The points raised include :—(1) the conditions in the dry season in respect to the 
welfare of the fly ; (2) the effect of grass burning on the numbers of the fly ; (3) the 
value of mopani country to the fly ; (4) the question of migration under the influence 
of hunger or other stimulus ; (5) the question of fly moving about infested areas with 
game; (6) the distance at which the fly can detect its hosts; (7) the following 
distance of female flies. 

In the following pages it is desired to emphasise the importance of these points, 
stating the writer’s own attitude of mind towards them in the light of available 
evidence, but without preferring any claim to having reached a final conclusion. 
Direct investigation of several of the points is planned for early attention if cir- 
cumstances permit, and it is hoped that other investigators will carry out similar 
researches so that definite solutions may be reached. 

(687) B2 


114 RUPERT W. JACK. 


Conditions in the dry Season. 


Dr. H. Lyndhurst Duke, of Uganda, in a recent paper* refers to the dry season as 
a time of stress and reduced breeding rate for the fly in that territory. Lloyd’s 
records concerning Northern Rhodesia are quoted as parallel observations, although 
the latter’s statement to that effect applied only to the earlier part of his work in the 
Luangwa Valley (Bull. Ent. Res. 11, p. 234) and his later work on the plateau led to 
exactly the opposite conclusion, namely that breeding was practically confined to 
the warmer part of the dry season (Bull. Ent. Res. v, p. 58). Lamborn in Nyasaland 
has made several statements on this pomt. For instance, in his Third-Report on 
Glossina Investigations in Nyasaland (Bull. Ent. Res. vii, pp. 29-50) on page 30 we 
have the statement: ‘‘ The small number of living pupae is to be accounted for by 
the breeding being at its nunimum during the late dry season” (July). On page 36 he 
states: ‘‘ Now as the dry season draws to a close pupae are being produced more 
freely;’’ and on page 46, “ The breeding season for G. morsitans is in full swing, just 
as the rains are about due.” It would seem that the period when the dry season 
“‘ draws to a close”’ and “ the rains are about due” might without inaccuracy be 
described as “ the late dry season ”’ in preference to July. Nevertheless Lamborn’s 
contention is quite clear, namely that the breeding season is at its minimum in July, 
and is in full swing in October, when the first rains may be anticipated both in 
Northern and Southern Rhodesia. In Southern Rhodesia, however, the October 
rains are normally very local and may not occur at all, and dry season conditions 
frequently prevail well into November. October and November (before heavy rains 
set in) are well known as the hottest months of the year and the term “ latter part of 
the dry season ”’ would appear naturally to include the period from the beginning of 
August to the opening of the heavy “ planting ” rains, which usually commence about 
mid-November. In some seasons they are, however, postponed until well into 
December, and in such circumstances, as the veld still maintains a dry season 
character, observations on breeding rates, etc., are still influenced by late dry season 
conditions, and this should be made clear. | 


Observations on the seasonal breeding rate in Southern Rhodesia are insignificant 
compared with those in Northern Rhodesia and Nyasaland, owing to the fact that 
no officer has been available to devote even the bulk of his time to the tsetse-fly 
problem. There is not the least doubt, however, that the fly may, under favourable 
conditions, breed very freely during the latter and warmer part of the dry season 
(August to November). Observations at Sipane Vlei, already published, leave no 
doubt upon that point. It would seem therefore that conditions in Nyasaland, and 
Northern and Southern Rhodesia are similar in this respect, at least as regards the 
plateaux. In Uganda on the other hand the published records of rainfall in the 
Masindi area and at Mpumu show that the climate is very different, the dry season 

being much less clearly marked and of much shorter duration. 
~ _In 1911 the writer called attention to a marked difference in the distribution of fly 
in the dry and wet seasons.f The dry season haunts of the fly in Southern Rhodesia 
have been described more than once. They consist of centres where shady evergreen 


*Bull. Ent. Res. x, p. 20, lines 11-13, and elsewhere. 
Bull. Ent. Res.‘ii, pp. 357-361. 


NOTES ON THE BIONOMICS OF GLOSSINA MORSITANS. 115 


trees exist and apparently correspond more or less to the areas termed “‘ foci,” 
“primary centres,” etc., described later in Central and East Africa. These dry 
season haunts undoubtedly vary to a considerable extent in their efficacy as favour- 
able habitats for the fly, as is indicated by the fact that some of them are intensely 
and others little infested. The type of dry season haunt where the fly is generally 
met with in the greatest numbers consists of a vlei, with the necessary evergreen 
trees along at least some portion of its margin, in a part of the fly area where vleis 
are scarce. Surface water may be present, or not, but the underground supply is 
always much nearer the surface than in the surrounding country, with the result that 
green grass starts growing with the warm weather long before the rains commence. 
The green grass in the vlei, especially after the dry grass has been burnt off and the 
young grass has begun to grow, is intensely attractive to grass-feeding animals, 
which often concentrate in considerable numbers around an isolated spot of this 
nature and move in regularly during the afternoon to feed, passing perforce through 
the haunts of the fly in doing so. In this way the flies obtain more or less regular 
meals and the whole conditions appear to be ideal for rapid increase during the latter 
part of the dry season, namely, regular food, warm weather and shelter for the 
larvae and pupae. Such centres might conveniently be termed “first grade foci.” 

In other parts of the fly areas rivers and vleis are much more prevalent, and intense 
concentration of both game and fly does not occur. The dry season haunts of the 
fly correspond, however, with the best grazing grounds at this time of the year, on 
account of the fact that the superior underground water supply supports both ever- 
green trees that shelter the fly and green grass that attracts the game. The term 
“second grade foci’ might be permitted in this connection. 

In other places again, the fly affects shady forest bordering rivers, streams or dry 
water-courses, and although not the main grounds, game has a tendency either to lie 
up in such situations during the day, or to pass through for the purpose of drinking. 
In the writer’s experience, wart-hog, water-buck, kudu, impala and small buck are 
very commonly found in this type of forest during the daytime, whereas certain 
types constitute the permanent home of bushbuck. Such spots are rarely heavily 
infested and may be termed “ third grade foci.” 


It must be borne in mind that animals of all sorts seek the shade during the heat 
of the day in hot weather, and as the fly is also dependent upon shade, fly and game 
must in general tend to meet at the time of year when shade is restricted more readily 
than at other times when shade, water and grass are everywhere to be found. 
Provided, therefore, that the temperature is suitable for breeding to proceed at the 
maximum rate, the dry season should in a general way be more favourable to the fly, 
assuming its dependence on the larger mammals for its food supply, than the wet. 

It does not follow, however, that this is necessarily the case in particular instances, 
Certain tracts of country dry up so thoroughly towards the end of the dry season 
that they are deserted altogether by most species of game. As has been remarked 
by several observers, however, wart-hog and duyker are usually to be found even 
after the grass has been burnt off over a wide area. These species, especially wart- 
hog, may serve to keep the flies from starving, but conditions cannot be said to be 
favourable to increase. Fly under such conditions is almost invariably very hungry, 
the proportion of the sexes caught tends to approach equality and pupae are 


116 RUPERT W. JACK. 


extremely scarce.* Fly appears to be extremely numerous and is certainly extremely 
pertinacious. Half-a-dozen flies that are eager for blood give a far greater impression 
of numbers than half-a-dozen “following” males. Fifty or sixty really hungry flies 
would give the impression of hundreds. Several investigators have indeed drawn 
attention to the fact that the numbers of tsetse under such conditions may be more 
apparent than real. . The writer can only speak with experience of one locality of 
this nature (Gorai River, Lomagundi), but a noteworthy point was the extreme 
localisation of the fly, few being met with except at the point where the path crossed 
the water-course, and here they seemed to swarm. When the writer camped at this 
spot for several days, the flies became very much less troublesome. Many pre- 
sumably fed on the two dogs accompanying the party, and a few on the natives. 
Some 80 odd were caught, and the party was then left in comparative peace. No- 
where else in the dried-up triangle of country formed by the Hunyani and Ambi 
Rivers with the escarpment were flies met with in any numbers. 

The following April no fly was encountered at the river crossing, but they were 
present in the neighbouring mopani forest in very moderate numbers, giving the 
impression of a decrease since the previous November. In regions such as this it is 
judged that the wet season must constitute the main breeding period, owing to the 
return of game with the rains, but that under these conditions the fly does not attain 
the same numbers as in areas where more favourable winter haunts are found. The 
visitor in the dry season meets, however, a large proportion of the fly population at 
once, whereas if game is plentiful he probably meets only a fraction. 

This argument may not apply universally to localities where fly seems numerous 
and game very scarce. In the area referred to it was actually shown that game was 
moderately plentiful when the country was not dried up. Data for times of year 
other than the dry season appear to be lacking in respect to most of the classic 
instances where fly apparently abounded in the absence or extreme scarcity of game. 

During the earlier part of the dry season, in May, June and July, the coldest 
weather occurs, and there is little doubt that the fly breeds less freely at this time of 
year and that the pupal period is more prolonged, both of which facts tend to check 
increase. On the whole, the fly tends to be less numerous at this time of year, 
particularly in July, but durmg August and onwards to the advent of the rains con- 
siderable increase occurs. With the advent of heavy and persistent rains the fly 
scatters and therefore appears to decrease suddenly, although this is probably only 
apparent. 

The answer to the question as to whether the dry season is in general a period of 
stress for the fly or not, except in so far as the breeding is influenced by temperature, 
would appear therefore to be in the negative as far as conditions similar to those of 
this territory are concerned, and the writer is of opinion that, excepting certain 
localities, far from being a time of stress, the latter part of the dry season is probably 
the most favourable and important period of the year under normal conditions. 
Furthermore, it would appear that the later the rains are in commencing, the greater 
the insects’ capacity for increase. That such conditions are not inimical to the fly 


*The writer is largely repeating results obtained by others in Central and East Africa, 
although observations in 8. Rhodesia are entirely in accord. 


NOTES ON THE BIONOMICS OF GLOSSINA MORSITANS. 117 


is shown by the fact that the latest onset of the rains recorded in recent years in the 
territory, namely in 1912, when no appreciable rain fell till towards the middle of 
December, was followed next year by the largest extension of fly recorded in connec- 
tion with the Sebungwe fly area. 


The Effect of Grass Fires. 

Another point is the stress laid by certain investigators on the efficacy of grass 
fires in reducing the fly. The present writer has been unable to obtain the slightest 
evidence that grass fires in this territory have any material effect on the number of 
fly, indeed the evidence to the contrary is almost conclusive. One of the most 
striking instances was recorded on the Gaori River below the escarpment in the 
Lomagundi district. Fly in November 1910 was still confined to a dry season haunt, 
which consisted in this case of shady trees on the bank of the dry water-course 
mentioned above. For some reason the grass had not been burnt until three days 
(according to the natives) before the writer’s visit, and the presence of still smoul- 
dering trees and the general appearance of the ashes of the grass showed it to have 
been burnt extremely recently. The fire had swept through the country over a very 
wide area. Fly was present in considerable numbers, as already mentioned, attacking 
the party very persistently. When a second visit was made to the spot in April of 
the next year, it was found that the grass throughout the “ focus ”’ stood as high as a 
man’s chin and was so thick that it was far too laborious to attempt to make headway 
through it. The fire the previous November must therefore have thrown up flames 
twelve to fifteen feet into the air. These fires are an annual occurrence, here as 
‘ elsewhere, and yet this little water-course is well known to all using the path which 
crosses it, as a fly centre, where the pest is always very much in evidence in the dry 
season. This was, as a matter of fact, the reason why that particular foot-path was 
followed. Many other less striking instances have come to the writer’s notice during 
the past ten years, and the accumulated evidence has induced an entire loss of faith 
in the efficacy of grass fires in reducing the fly. As a matter of fact if tsetse-flies 
could not avoid grass fires they would surely long ago have been exterminated in 
parts of the country inhabited by natives. Of course even in Southern Rhodesia 
much patchy burning of the grass occurs, but the long, intensely dry winter pre- 
sumably favours burning over wide areas more than conditions in Uganda, for instance, 
It is difficult to see how the flies could avoid a roaring furnace like that along the 
Gorai River by mounting in the air unless they went to a very great height (for a fly), 
but the contiguous mopani would afford a refuge, as in this type of forest the grass is 
always very short, and is sometimes almost altogether wanting. If large areas of 
country infested with fly were covered with long thick grass better results might be 
anticipated. Possibly this is the reason that greater effect on the fly has been 
noticed elsewhere. 


The Value of Mopani Country to the Fly. 

Both Duke and Fiske, referring to Uganda, speak of the fly showing hunger in the 
long grass season, which is of course the latter part of the wet season and earlier part 
ofthedry. This is attributed to difficulty in locating game. The point is extremely 
interesting, because from Duke’s description of the Masindi area comparatively long 


118 RUPERT W. JACK. 


grass apparently occurs over wide areas. Conditions differ in various parts of the 
fly-infested areas of Southern Rhodesia, but in the great majority mopani belts are 
a feature of the country, and the fly certainly shows a strong preference for this type 
of forest in the wet season, deserting its dry season haunts where the grass is usually 
long and thick, sometimes very long. The sweet short grass of the mopani is very 
attractive to many species of game in the wet season, though the reverse in the dry, 
and the fly certainly has the best chance of meeting and perceiving game in this type 
of country during the rains. The mopani grass, however, dries up very quickly after 
the cessation of rains, being shallow-rooted and of little substance, so that it loses. 
its special attractiveness very shortly after the close of the wet season. Nevertheless, 
this type of forest is much haunted by impala and wart-hog even at that time, and 
many other species are commonly seen in it, if game is at all plentiful in the neigh- 
bourhood. Whilst, therefore, it is probable that m Southern Rhodesia the long- 
grass season is not in general a time of stress for the fly, the fact is m no way opposed 
to Fiske’s observations. In this connection it may be mentioned that the only sex 
count made by the writer in the long-grass season was in April 1911, with fly taken 
in mopani, and resulted in 53 males to 3 females, by far the most extreme disparity 
yet recorded in the territory. The fly’s habit of avoiding thick undergrowth would 
indeed suggest a difficulty in finding its hosts in such an environment. It is quite 
possible that in certain parts of the fly areas in Rhodesia a time of stress for the fly 
may occur in the long-grass season, as the mopani is not ubiquitous, being absent, so 
far as the writer is aware, from most of the Jetjenini fly area and from certain parts. 
of the Sebungwe area. Prospectors and hunters in this territory have constantly 
associated mopani and tsetse, and although obviously not essential to the tsetse, it is 
quite possibly of value to the fly under certain conditions. According to Fiske’s 
observations it ought theoretically to be of value in the long-grass season, and possibly 
it affords a refuge from grass fires. 


The Question of Migration. 


The next point is the question of the migration of fly, corresponding to the move- 
ments of game, or under stress of hunger. Now it is far from the writer’s intention 
to adopt a dogmatic attitude on this subject, but it would seem difficult to reconcile 
any habit of this nature with a number of known facts. First, in regard to the ques- 
tion of fly migrating with game, we have the phenomenon of restricted fly-areas and 
their mode of extension. It has already been pointed out (Bull. Ent. Res. x, p. 88) 
that the limits of a fly area are not necessarily permanent. The permanent limits are 
marked by the impingement of favourable on unfavourable country; that is to say, 
the country beyond the limit is for some reason or other unsuited to the tsetse. 
Transitory limits are, however, formed by the high-water-mark, so to speak, of the 
advancing flood when the fly is spreading, as has been the case in this territory since 
the rinderpest. Transitory limits would also be apparent if the pest were receding. 


The advance of the pest is, however, comparatively slow, and in no way com- 
parable with the powers of movement of the fly itself or of game. Much potential 
fly area, known to have been infested in pre-rinderpest days, is still free from the 
pest, although the latter is gradually occupying more and more of its old country. 


NOTES ON THE BIONOMICS OF GLOSSINA MORSITANS. 119 


Is it possible for these conditions to obtain if the fly had a habit of migrating with 
herds of game, apart, of course, from the males’ habit of temporarily accompanying 
moving animals and human beings? Herds of certain species of game are more 
or less constantly on the move in and out of the fly area, and if the fly accompanied 
them in their wanderings the whole of the potential fly area would certainly be very 
quickly occupied. 


> 


It may be contended, however, that the fly only follows “ trekking ” game under 
stress of hunger. This appears to imply a rather marvellous instinct on the part 
of the fly. Any fly suffering from hunger that encounters a herd of game is likely 
to be a full-fed fly in a space of time measured by minutes, and is then not in a 
condition to follow anything for some time. Must we, therefore, suppose that the 
fly restrains its appetite in order to accompany the herd into another part of the 
country ? Again, game travels as a rule by night, and though there are numerous 
records of fly “‘ biting ” on warm nights, it is essentially of diurnal habit. Finally, 
we have apparently no records to show that the female flies follow animals or human 
beings for any appreciable distance at all, whether they are hungry ornot. The 
writer’s observations are all to the contrary, and published accounts apparently 
give no record of other than males exhibiting an obvious following habit. Much 
argument has in fact been based upon the supposition that the female flies only seek 
animals and human beings for the purpose of feeding, whilst the males follow for 
other reasons. 


These considerations have led the writer to be extremely sceptical concerning 
the fly’s alleged habit of migrating with game. Regular movements of game in a 
given direction might, it is admitted, help the spread of the fly in that direction. 
The allusion is to the daily movement of game from its feeding grounds to water, 
as, for instance, game which feeds in the neighbourhood of a dry river and moves 
across to a flowing one to drink (e.g., the Mzola and Kana Rivers in this territory). 
In the same way, regular traffic along a road would tend to spread the fly, if condi- 
tions admit of spread, but this is quite apart from the question of fly migrating 
with migrating game. 


The second aspect of the question is that of fly migrating without help from 
moving game, when pressed by hunger. This would be a serious state of affairs ; 
as they might be attracted by the settler’s herds of cattle, take up their residence 
in the vicinity of his “ kraal” and deposit their larvae under his orange trees. 
Such a development might well have happened in the writer’s experience if hungry 
fly had a habit of migrating even three or four miles to a food supply. The important 
point is that it did not happen. The idea of this form of migration has, as a matter 
of fact, always puzzled the writer. How is it supposed to take place? Do the 
tsetse send out scouts like bees to locate a better piece of country and then rise in a 
swarm to follow these guides to the new locality 2? Does some instinct inform 
them that better conditions prevail elsewhere, so that they rise with a common 
impulse and migrate thither? Do they, under stress of hunger, migrate en masse 
at random guided by chance, or perhaps the direction of the wind? Do they migrate 
by a series of short journeys, gradually tending day after day in one direction ? 
Unless the movement takes place at random, this presupposes a knowledge of where 


120 RUPERT W. JACK. 


better feeding grounds are to be found. Or, finally. do they migrate individually ? 
In the latter case they might take many directions and a corresponding increase 
in another part would not be likely to accompany the decrease in the original 
centre. Fortunately for the neighbouring settlers, where we have had a clear case 
of hunger conditions as regards game, in the Hartley district, the diminishing fly 
became gradually restricted in range as the game decreased, and still adhered to 
their old haunts, whilst numerous cattle worked with more or less impunity amongst 
the Shagari mines a few miles away on the one side, and settlement proceeded 
along the railway line with rapidly decreasing losses on the other. The destruction 
of the usual haunts by the felling of the forest did, however, appear to cause a scat- 
tering of the tsetse, which died out, probably in consequence. The fly along the 
Gorai River, mentioned above, were obviously very hungry at the time of the writer’s 
visit in November 1910 (out of some 82 flies caught actually more were females 
than males), and from all accounts this is an annual occurrence at this spot; but 
although game was more abundant on the Hunyani a few miles away, the flies 
apparently preferred to endure hunger in this area rather than migrate. They 
were not, it must be confessed, absolutely starving, as wart-hog and duiker were 
actually seen in the fly haunts; but their meals were without doubt sufficiently 
irregular,to keep the appetite of the majority very keen. Fly, on the other hand, 
was comparatively scarce on the Hunyani where more game was to be found. 


The writer has, in fact, been able to find no evidence during the past ten years 
in this territory that the fly has any tendency to migrate, apart from the seasonal 
scattering during the wet season, as a result of which the pest is much more uni- 
formly distributed over the infested area than during the dry season. Although 
new areas have been invaded by its gradual spread, no incursion of fly into localities 
other than those immediately adjacent to the former infested areas have occurred. 
It is readily admitted that an apparent migration might take place in the course 
of time, the fly increasing in a newly invaded area and decreasing in the old, but this 
does not imply migration. In any case in Southern Rhodesia the invasion of new 
areas, in the writer’s experience, has not been accompanied by any corresponding 
reduction elsewhere, nor has reduction in an old area been accompanied by any 
noticeable increase elsewhere. The fly appears to increase or decrease in any area 
solely in relation to its ability to breed successfully. 


On the whole there seems little reason at present to believe in any form of 
migration in respect to Glossina morsitans, other than the seasonal scattering with 
the advent of the wet season, and forced movements induced by destruction of the 
forest. In the latter case the fly naturally follows the receding shade. 


The Question of Fly moving about Infested Areas with Game. 

The idea that fly locates game and follows its movements more or less continuously, 
at least within the limits of infested country, is not absent from the writings of 
serious investigators. In this connection it is essential to distinguish between 
following for a limited distance, as is the well-known habit of the males, and possibly 
of the females to a much less degree, and the alleged habit of the fly attaching 
itself to the herds and accompanying them in their wanderings day after day. The 
writer has been able to find no direct evidence adduced in support of this idea, 


NOTES ON THE BIONOMICS OF GLOSSINA MORSITANS. 12] 


and there are certainly many reasons why such a habit is, to say the least of it, 
unlikely. The question is intimately connected with that of fly migrating with 
game ; for if the fly followed herds of game within the infested area, there appears 
to be no obstacle to its following the herds beyond it, provided that the country 
entered is potential fly country. This, as an attempt has been made to show, is 
at least not its general habit. The further difficulties in the way of accepting 
this idea, mostly already mentioned, may be briefly summed up as follows :— 
(1) female flies are apparently not known to follow to any great distance at any time ; 
(2) gravid females seek seclusion, and are not in the least likely to attempt to follow 
a moving herd ; (3) hungry flies tend to feed fully, and full-fed flies abandon their 
hosts, which are unlikely to remain in the vicinity until the flies recover; (4) the 
fly is diurnal in habit, whilst game moves largely at night. It would seem therefore 
that the only individuals capable of following game even for a few hours are those 
which do not desire to feed. The majority of observations indicate that only the 
males exhibit a tendency to follow, although the possibility of non-gravid females 
doing so is not altogether excluded. In any case it is well known that even the 
males do not follow human beings for more than an hour or so, but gradually fall 
away, and there seems little reason to think that they would exhibit a different 
habit in respect to game. Even supposing they followed till nightfall, they must 
surely lose touch with the herd after dark. 

It appears probable that the tsetse-fly neither ranges the forest in search of its 
prey, nor follows it when encountered for any great length of time, but that it waits 
for the animal to come within the range of its perceptive powers. The fact that 
fly is constantly met with at the same spots* strongly suggests ambush rather than 
pursuit, and the writer has certainly no experience of having encountered fly in 
circumstances that suggested anything else. 


The Maximum Distance at which the Fly can detect its Hosts. 

Direct experiments are needed and planned to determine this point as far as possible. 
It appears necessary to distinguish between the maximum distance at which the fly 
readily perceives its prey, so that a man or animal would not pass by without being 
observed, and the distance at which it might locate say a herd of game grazing for 
several hours in one spot. 

Some of the early explorers and hunters who encountered tsetse-fly were most 
emphatic that a very short distance might separate complete safety from certain 
death for their cattle, a narrow stream being sometimes mentioned as the dividing 
interval. Like other statements emanating from untrained observers this assertion, 
though undoubtedly greatly exaggerated, appears to have some foundation in fact. 
It would be possible to mention numerous personal observations of having been very 


*The writer is aware that observations have been published of fly having been 
encountered in quantity at certain spots on one occasion and only in very small numbers 
at the next visit. During the wet season this is quite likely, as the fly is not confined 
by lack of shade, and when carried even for a short distance is hardly likely to return to 
. exactly the same spot. Even in the dry season the passage of a herd of game might 
make a very material difference for a few hours, the hungry flies feeding and seeking 
seclusion, and the non-hungry males following the herd. Nevertheless, allowing for the 
difference in distribution in the wet and dry seasons, the statement is substantially 
correct. 


123 RUPERT W. JACK. 


little molested by fly on one side of a vlei, whereas on crossing to the other, some two 
hundred yards or so away, the party was immediately assailed by great numbers. A 
rather striking instance of the fly’s limitations in this respect occurred in October 
1919. The writer passed along a foot-path about 9 a.m. to examine a salt vlei near 
the Shangani for fly. None was seen till this vlei was reached and an hour was spent 
walking all round the margin, during which time only four tsetse were caught. 
Returning, the route taken aimed to cut the path at an acute angle, and when within 
100 yards of the path on reaching a shady tree a number of tsetse suddenly attacked 
the party and “ bit ” with extreme voracity. Yet about an hour earlier the writer 
with his natives had passed within a hundred yards without being perceived. It. 
cannot be proved definitely that the fly were there an hour earlier, but there was no 
spoor to indicate that any game had passed in the interim, and as the morning was. 
hot the fly would hardly have crossed the leafless mopani to this particular tree. The 
writer was extremely impressed by this particular instance, and as a matter of fact the 
limitations in the fly’s capacity for readily detecting a food supply has been a constant 
source of surprise throughout investigations now extending over ten years. A move- 
ment of fifty yards or so in a fly haunt frequently brings an accession in numbers of 
hungry insects and in numerous cases when all the flies seen were being collected, 
this phenomenon has been particularly marked. If it be accepted as an axiom that: 
an animal or human being attracts all the hungry tsetse within a radius not exceeding 
the fly’s powers of perception, then the limitations of the fly in this respect have been 
proved again and again; for fresh hungry flies have undoubtedly been constantly 
encountered after a very short movement, whilst some two hundred yards or so has. 
been found sufficient to keep a party comparatively free from attack whilst the fly 
swarmed that distance away. To mention another instance, in September 1913, 
the writer halted for breakfast on the south-east side of Sipane Vlei, between the 
Sengwa and Sassame Rivers in the Sebungwe district. A certain number of tsetse 
had been picked up en route, and the boys fetching water brought in more, so that a 
few bites were received. Some time after breakfast the writer took three natives to 
the north-west side of the vlei; some 300 yards away, and found the fly in numbers, 
some 87 being caught with one net in an hour. The wind was from the south-east 
and had been blowing over the party towards the fly haunts for at least two hours ; 
yet the bulk of the flies had not been attracted over this comparatively short distance, 
though they attacked the party in considerable numbers when their haunts were 
entered. Further visits to this spot have been made since, and it has been found 
that by camping on the south-east side of the vlei the attentions of fly could always 
in a great measure be avoided. 

From the foregoing considerations it is believed that the tsetse-fly is only capable 
of readily detecting its hosts at a comparatively short distance, probably less than a 
hundred yards. It might, however, be longer in the wet season, when scent pre- 
sumably carries further than in the dry. 

If this limitation be proved, the fact increases the difficulty of accounting for the 
presence of fly in numbers in areas where large mammals are markedly scarce. It 
apparently remains to be proved, however, that fly occurs in numbers in any tract of 
country where large mammals are markedly scarce at all seasons of the year. If 
such a phenomenon does exist, the writer admits frankly that it appears irreconcilable 
with the theory of the vital association of the two forms of life. 


NOTES ON THE BIONOMICS OF GLOSSINA MORSITANS. 123 


Is it not possible that this limitation might account to some extent for the gre- 
garious habit of the tsetse, the flies benefiting by the combined perceptive powers 
of a number instead of relying on those of single individuals ? The value of numbers 
in regard to perception will hardly need urging upon anyone who has hunted antelope 
or other gregarious game. 


The Following Distance of Female Flies. 


It appears very important that attempts should be made to determine as accurately 
as possible the maximum distance to which female flies may be carried by game, 
man, etc. The point has a great bearing upon the question as to what would con- 
stitute an effective barrier to the fly’s advance, and also upon the question of the fly 
as a whole migrating with game or moving about the fly area in company with its 
hosts. The writer.has as yet been able to prove following on the part of the females 
up to only about 400 yards, although one instance has occurred where a specimen 
had apparently followed very much further than this, over a mile in fact. The ob- 
servation in the latter case was, however, liable to considerable error. Some 
experiments in this connection were planned for the writer’s visit to Sipane Vlei in 
August 1919, but could not be carried out owing to the unexpected scarcity of fly in 
that centre. 


Sipane Vlei and some other Localities in 1919. 


In a recent paper * the writer pointed out that the somewhat delicately balanced 
state of affairs which exists at the most favourable dry season haunts of the tsetse, 
namely the margin of vleis containing green grass in the dry season, is apparently 
hable to be somewhat easily disturbed by any agency tending to prevent game from 
visiting the vlei with its normal regularity. This disturbance, it was pointed out, 
would enhance the effect of game reduction by hunters, and might also have operated 
in the rinderpest epizootic, which undoubtedly induced abnormal movements of the 
panic-stricken remnants. These views have received some indirect support during 
the dry season of 1919, when for the first time since the rinderpest a reduction of fly 
in areas of the territory unaffected by civilisation has been recorded. This reduction 
is local, and extension since the previous year was recorded in one other part of the 
country, and may have occurred elsewhere. A reduction has been reported 
in the northern part of the Umniati fly area by Dr. Alec Mackenzie of Gatooma. 
Dr. Mackenzie’s statement is to the effect that he found no fly at all in December 1919 
in many parts where they are usually numerous near the Sakugwe and Umniati 
Rivers on the road from Gatooma to the Emerald Mine, and only a few finally at the 
headwaters of the Mvumvudzi and Urungwe Rivers, visited for the express purpose 
of locating the fly. There is, however, one spot where reduction can be proved on 
something like a mathematical basis, namely Sipane Vlei, lying between the Sengwa 
and Sassame Rivers in the Sebungwe district. 


The rapidly-expanding Sebungwe fly area reached the Sengwa River from the west 
by 1910, and fly was first recorded at Sipane in 1913, though it may have been present 
at least a year earlier. In late September of that year it was present in great numbers, 
and 87 were caught with a net in one hour, in order to establish some sort of basis for 


*Bull. Ent. Res. x, p. 83. 


124 RUPERT W. JACK. 


later observations. In November 1914, the flies in the vlei were “ like a swarm of 
bees ” around the individual who penetrated their haunts, and the ease with which 
pupae were collected during that visit has already been recorded. In August 1916, 
fly was extremely abundant, and living pupae could be found (some 30 were collected 
in a few hours), though the latter were not as abundant as in November 1914.* In 
late August 1919, the writer spent ten days at the vlei, for the purpose of chopping 
down the evergreen trees as an experiment. Hardly half-a-dozen fly were seen in 
the course of the day, where formerly they swarmed, and search for living pupae 
was entirely unsuccessful, notwithstanding that much more time was available than 
on previous visits. Only a few empty cases were found in the usual breeding places, 
and from these the flies had apparently emerged normally, there being no indications 
of parasitism. 

In seeking a probable explanation of the decrease it was impossible to ignore 
the marked influence that the heavy rainfall of the two previous wet seasons had 
had on the country. The wet season of 1917-18 was the heaviest recorded since 
1890-1, and was of a peculiar nature, the skies remaining overcast for weeks together, 
so that the maize crop suffered severely from excessive wet and lack of sunshine ; 
conditions were thus ideal for maximum penetration. The following season’s 
rainfall was also above the average. The first difference noted in the appearance 
of the country was the fact that the gusu (Brachystegia) forest bordering one side 
of the vlei had not yet lost its foliage, although quite leafless in August 1916 (see 
photograph, Bull. Ent. Res. x, p. 90, pl. u, fig. 2). The same remark applied 
in a lesser degree to the thorn thicket (isi-nanga). The mopani, on the other hand, 
was practically leafless, except half-a-dozen trees in a wet situation on the edge of 
the vlei itself. These were in full leaf, having apparently grown out again since 
the fall. The gusu, it may be remarked, showed no young foliage and the leaves 
were falling sufficiently rapidly to show an appreciable diminution of shade during 
the ten days of the writer’s stay. Careful inspection of the environment revealed 
a considerable extent of green grass to the south-west and west of the vlei, which 
was not in evidence during previous visits. Further, game was remarkably scarce 
in comparison with previous experience, and such as occurred was not feeding at the 
vlei itself, but on the green grass mentioned away from the vlei. With the exception 
of two small herds of impala, one kudu cow, one wart-hog, a rhinoceros, a wild 
dog and several duiker, no game was actually seen, in spite of daily excursions 
in all directions. Fresh spoor of a herd of zebra and another of sable was found 
in the vicinity. A lion drank one night at the vlei, as also did a solitary buffalo bull. 
This sounds a rather formidable list, but it represents the results of ten days explora- 
tion of the country within a radius of about eight miles of the vlei. A pack of 
wild dogs was apparently hunting the neighbourhood, and this may account to some 
extent for the scarcity of antelope, but the main point ascertained was that the few 
antelope present in the vicinity were not visiting the vlei for the purpose of feeding 
as is usually the case. They apparently found more attractive fare in the stretches 
of green grass away from the vlei (see sketch-map). 


*In connection with the comparative smallness of the numbers quoted here and 
elsewhere it should be noted that no trained natives were available and that the pupae and 
flies were practically all collected by the writer personally. 


NOTES ON THE BIONOMICS OF GLOSSINA MORSITANS. 125 


Further evidence in support of the theory lies in the fact that fly was actually 
found more readily, though not at all abundantly, in the green grass areas away 
from the vlei, where such game as occurred was grazing, than at the vlei itself. The 
general prevalence of shade apparently rendered this possible under the prevailing 
conditions, although this is not usually the case at this time of year. A few fly were 
even met with in the gusu itself, which was unattractive to the majority of game, 
but only three were seen in a walk of several hours. 


- 


enya 


Manyoni R. 


Scale 9miles to linch. 


The obvious question is whether the fly had decreased from failure of the birth 
rate to equal or exceed the death rate, or from the majority having migrated. There 
was certainly no corresponding increase of fly within a radius of seven or eight miles 
of the vlei, as the country within this radius was thoroughly explored at the time. 
Nor had they shifted to the nearest known dry season haunt, which is Matjagenya on 
the Gadzi River, some seven miles away. There is no evidence of the appearance 
of fly at any point in this region outside the limits of the known area, and if the fly 
had migrated en masse in this direction it must have been noticed, as the country 
is inhabited. As a matter of fact the natives questioned stated that they knew of 
no fly outside the previously marked limit in this locality. It would appear, there- 
fore that if the fly had migrated it must have moved back into the infested country 


126 RUPERT W. JACK. 


and for a distance exceeding the radius explored around Sipane Vlei. It is obviously 
possible that this may have occurred, but it appears very unlikely. Whatever 
be the correct explanation, it is certain that Sipane Vlei had ceased for the time 
being to serve the purpose it had served in the past, as a favourable breeding haunt 
for the fly during the latter part of the dry season. During the season of dispersion 
a considerable area of country would be affected by this fact. 


It is interesting to note that some years ago an experiment was planned in connec- 
tion with Sipane Vlei with a view to determining whether the fly there was dependent 
upon the visits of big game or otherwise. The plan was to surround the whole vlei 
with a game-proof fence and thus keep the game away during the season of concen- 
tration. It was not carried into effect on account of the doubtful practicability 
of effective control at such a distance from communications. It would almost 
seem, however, as if meteorological conditions, in reducing the peculiar attraction 
of the vlei to game, had effected a demonstration instead. Unfortunately further 
observations bearing on this point will not be possible at this centre as the ever- 
green trees on the margin of the vle1 have now been cut down. 


Time was not available for a detailed inspection of other dry season haunts in the 
same region, but fly was scarce at the crossing of the Gadzi River, known to the 
natives as Matjagenya, on the way to Sipane, though formerly numerous. The 
natives resident at this point also bore out this observation. Again, at Depota 
Vlei some seven miles east of Matjagenya, no fly at all were seen, and some natives 
who had taken up their residence there the previous year had a number of apparently 
healthy dogs. They stated that they never saw fly near the vlei, although a very 
few were seen there by the writer in 1916. Winter shade is scarce at Depota, other- 
wise it would be a typical first-grade focus, as it is attractive to game as a rule. 
The writer has in the past seen eland, kudu, sable, impala, zebra, and sessaby in the 
vlei, and the fresh tracks of buffalo and rhinoceros in addition. In August 1919 
only one duiker was seen in the neighbouring forest, and the tracks of three kudu. 
For the moment, therefore, the fly seems to be retrogressive rather than otherwise 
in this region, and it is essentially a region of isolated dry season haunts. 


On the other hand, at the point where Advance has been recorded definitely, 
namely up the Shangani River, the region is one of diffused dry season haunts 
(second and third grade foci), and game was very plentiful in October 1919, though 
their numbers had doubtless been augmented owing to the organised shooting 
further down the river which commenced in June. Here then we have an illustration 
of “‘ the fly following the game,” only as the area of increase is some ninety miles, 
as the crow flies, from Sipane Vlei the fly can hardly have migrated thither. The 
presumption is that conditions the previous year favoured peal in the one area, 
and were unfavourable in the other. 


Whilst conditions in the northern part of the Umniati fly area have no doubt 
been affected to some extent by the efforts of hunters in shooting game for the 
“Emerald ” and “‘ Copper Queen ” Mines, and the portion to the east of the river 
considerably affected on account of being in the open shooting area, the writer 
has been, so far as is known, the only European visitor to Sipane Vlei for several 
years past. 


NOTES ON THE BIONOMICS OF GLOSSINA MORSITANS. 127. 


It is not claimed that the only way in which one or two abnormally heavy wet 
seasons could affect the tsetse-fly lies in inducing a wider distribution of water and 
attractive grazing during the dry season. The involved interrelationship between 
the tsetse fly, its diseases and enemies, the enemies of its enemies, and so on, is 
almost a closed book at present, and this interrelationship is no doubt affected in 
some degree by the season. It is only intended to point out a possible explanation 
of the observed phenomenon as a basis for future research. The abnormally heavy 
tains of the past wet season have certainly affected the appearance of some parts 
of the country in the dry season to a marked degree, and the fly has undoubtedly 
decreased in certain areas, one of which at least, as the writer has endeavoured to 
show, has also been affected by the increased rainfall. | 


= 
@ 
1°) 
~ 


me 6 


Sketch Plan of 
Sipane Vlei and 
Neighbourhood. Aug. 1919. 


E; vergreen trees 
indicated by black dots. 


Scale about 1-2 m. to theinch. 


Curiosity naturally suggested reference to the records of rainfall in the territory 
with a view to ascertaining whether any record of one or two seasons of exceptional 
severity preceded any definite information concerning reduction of fly. The 
available rainfall statistics unfortunately date no further back than 1888 and in the 
(687) o 


128 RUPERT W. JACK. 


intervening period there is only one record of two consecutive wet seasons comparable 
with those of 1917-18 and 1918-19. These are the seasons 1889-90 and 1890-1. 
The total fall in these two seasons at Bulawayo considerably surpassed that of the 
recent two seasons, the number of inches recorded being 87°81 in the earlier couplet 
against 76°13 for the later. Was this heavy fall followed shortly by any marked 
reduction of tsetse? There is some suggestion to the effect that it was, at least in 
certain parts of the territory, although it is of such an indirect and unsatisfactory 
nature that it is practically valueless. 

It is curious, however, that during this very time, a marked recovery from a 
period of decrease seems to have taken place in the region of the Zambesi. In a 
foot-note to a paper by the writer (Bull. Ent. Res. v, p. 100), Captain Selous’ 
opinion is recorded that by 1888 “ the belt shown to the west of the Victoria Falls 
had disappeared,” and “ that much of the fly to the east of the Falls had also gone 
by that date.’ This statement was doubtless substantially true in 1888, in fact 
there is other evidence that fly was decreasing in this area during the preceding 
years (v. Dr. Holub, Austen’s Monograph, p. 203). Nevertheless Selous’ statement 
is misleading and should be corrected in case other investigators may use it as a 
basis for argument. The writer has first-hand information to the effect that fly 
‘was numerous between the Falls and the Zambesi-Chobe confluence in 1893, and 
it is apparent that an increase of fly took place subsequent to 1888. The first-hand 
evidence referred to is contained in a report by Mr. A. Giese, Cattle Inspector, to 
the Chief Veterinary Surgeon, Salisbury, dated 9th July 1918. The report is 
worth quoting in full, although some of the matter does not bear on the present 
argument. It appears that the fly did actually disappear from this part of the 
country at the time of the rinderpest, whatever may have been the position of 
affairs near the Limpopo. 

Mr. Giese’s report is as follows :— 

“ 9,vii.1918. The following notes are reminiscently penned because they may 
‘serve aS a guide as to where to expect the tsetse-fly, now re-appearing west of the 
‘Gwaai,* to spread to should this re-appearance be only a return to localities driven 
out from or killed by the rinderpest, and not as well to localities which the fly had 
already voluntarily disappeared from before the rinderpest arrived. 

“Tn 1861-2 Baldwin(?) brought the first waggons to Deka commencing the track 
which eventually grew into what is now known as the Pandamatenga or western border 
road. He had to leave his wagons at Deka and proceed to the Zambesi on foot as the 
intervening country was infected by fly. 

‘“ Somewhere about the middle seventies Geo. Westbeach established his later far- 
known trading station at Pandamatenga, when the fly had receded further north 
somewhere north of Gazume Vlei. At the same time, or a little later, transport could 
be taken to the Victoria Falls and to Bingua Spruit—about railway cottage 277—from 
where Westbeach’s waggons used to fetch grain traded in the fly-infested country 
to the east ; also that broken stony country north of the present Matetsi siding was 
believed to be free from fly right up to Victoria Falls. 


* No fly has as yet been found west of the Gwaai, but outbreaks of trypanosomiasis 
have occurred during the past three wet seasons. 


NOTES ON THE BIONOMICGS OF GLOSSINA MORSITANS. 129 


“In July 1893, the writer’s first appearance in these parts, ox transport was left at 
Gazuma, but fly was found between Leshuma and Kazungula, Chobe and Zambesi 
junction, and from Kazungularight up the Chobe and right down to within a few 
miles of the Victoria Falls, fly being particularly aggressive on the latter stretch. 

“Tn 1894-5 during various trips to what is now the Wankie Colliery all animals 
had to be left at Deka; in fact when the writer pegged the present Deka farm in 
1895 animals were not allowed near its 8. E. beacon. 

“ During these latter years the southern boundary of the fly area ran roughly from 
Galobe—Gwaai junction more or less west but north of that open park-like country 
round and to the south of Chumagu Malishi and Makolo to places near the present 
Kennedy’s Halt, and hunters used to take their ox waggons thus far north. 

“* At the end of 1897 the writer took a horse up the Chobe and in 1898 six spans of 
donkeys to the present Wankie Coal Mine, all fly having disappeared then.” 

In answer to direct queries by the Chief Veterinary Surgeon Mr. Giese replied as 
follows under date 30.vii.1918 :— 

1. “ What species.died of rinderpest ? 

“Eland, buffalo and kudu principally, and in the order mentioned as regards 
numbers found dead and the occurrence of the species at the time. The greater 
mortality among the eland was not due to their being more susceptible to disease but 
to their occurring in much greater numbers than the other game. This they still do. 
Of other game reedbuck probably suffered most. 

2. “ What was the rate of mortality ; were there any survivors % 

“In 1898 the only surviving buffalo, between Kazungula and the Gwaai were one 
bull, 3 cows and 3 calves, running 7 miles west of the present Colliery, and a somewhat 
similar number on the lower Inyantue ; which means that these were the only buffalo 
left in Matabeleland west of the Gwaai, because their habitat did coincide with the 
tsetse-fly. 

“What the number of buffalo was in pre-rinderpest days one cannot say, as they 
were scattered owing to the natives carrying guns ; but the increase of these 7 animals 
(survivors) is a fairly compact herd of some seventy animals now confining themselves 
to a belt of country 20 miles by 30 miles to the west of the railway. The Inyantue 
herd is a little larger. 

“There were stray survivors of kudu and they also have more than recovered 
owing to the absence of firearms; there are more kudu than any other antelope, 
Sharpe’s steinbuck excepted, in the broken and hilly part of the Wankie District, 


3. ““ Did the fly disappear at once ? 

“ The writer left Deka on 15th January, 1896, when fly were as stated in his notes, 
arrived in Bulawayo mid-February and was sent for from the C. C’s office to give 
an account of what he knew of rinderpest which was then approaching. When he 
returned after the Rebellion in 1897 the fly had disappeared.” 

The record is extremely interesting in showing that fluctuations in the distribution 
of fly on a moderately extensive scale have occurred in the past apart apparently 
from overwhelming game destruction, although the hunters appear to have been of 
opinion that the decrease of fly during the period preceding 1888 was due to their 

(687) o2 


130 RUPERT W. JACK. 


inroads into the number of buffalo. As a matter of fact, events in this region seem 
to have contributed largely to the solidifying of Selous’ opinion that the fly was 
dependent upon buffalo. 

The rainfall in the Zambesi valley frequently differs very considerably from that 
in the remainder of the territory, and it is unfortunate that no records in this area _ 
for the period concerned appear to be available. 


The records, are however, instructive in regard to the influence of a series of seasons 
of low rainfall. At Kariyangwe in the Sebungwe district the rainfall recorded for 
the three seasons 1909-10, 1910-11 and 1911-12 was extremely low. This station 
was then closed down, but referring to the Wankie records, somewhat further from 
the scene, we find that the next two years were also very low indeed. During this 
period the extension of the Sebungwe fly area proceeded at its maximum speed, as 
may be seen by a glance at the author’s map (Bull. Ent. Res. x, map 1). The 
records of advance in this area are moderately accurate as regards the three year 
intervals marked on the map, with the exception of the north, where the discovery 
of tsetse-fly on the lower Sengwe River in 1913 indicated that the previous limits in 
this area, visited only at long intervals by officials who traversed the country rapidly, 
were probably faulty. It is not possible, however, to obtain accurate yearly records. 
The dry season of 1913 was remarkable for advance in almost every direction, ex- 
cluding a portion of the western limit where the fly appears to have reached the 
division between favourable and unfavourable country. The rains were very late 
in 1912, no serious showers falling until near the mjddle of December. Dry season 
conditions were thus unusually prolonged, and it is believed that such conditions 
favour increase of fly. 

To recapitulate, whilst the writer is far from attaching any exaggerated importance 
to these limited records, they are certainly suggestive of abnormally heavy wet 
seasons having a deleterious effect on the fly in certain areas, and of a series of seasons. 
of low rainfall, especially if late, favouring increase. Intense and prolonged dry 
season conditions, such as occur during a cycle of years of low rainfall, are judged to 
favour the fly at centres which resist the action of drought best, owing to the con- 
sequent concentration of game. A rise in the water-table after an exceptionally 
heavy season or two causes a disturbance in the usual conditions by inducing a more 
general distribution of good grazing and water throughout the country, and a pro- 
portionately reduced attraction of the “ focus ” to game. 


It is much to be regretted that the writer’s itinerary in 1918 did not include Sipane 
Vlei. The tremendous rains of the 1917-18 season had a very marked effect on the 
forest lying between the railway line and the Gwaai River on the western side of the 
territory, and it is reasonable to suppose that conditions at Sipane Vlei may have 
been similar to those in 1919, seeing that at Gokwe, the nearest meteorological station, 
43°03 inches fell in the 1917-18 season and 33°10 inches the 1918-19 season, although 
even the latter is well above the normal for that district. If the writer’s theory is 
correct, fly should have been numerous and hungry in the dry season of 1918, pupae 
being, however, scarce, resulting in a great diminution of fly by 1919. 


This theory is very much in accordance with observations published by Lloyd 
(Bull. Ent. Res. vii, pp. 67-79) with reference to Northern Rhodesia. Lloyd draws 


NOTES ON THE BIONOMICS OF GLOSSINA MORSITANS. 131 


a most necessary distinction between apparent and real abundance of fly, and shows 
how the percentage of females caught in the ordinary way varies in inverse ratio, 
and the number of pupae secured in direct ratio, with the abundance of game. Here 
we have direct observations indicating that scarcity of game induces hunger and 
hunger inhibits breeding. Both these observations have received support from 
other investigators, including the comparatively scanty observations in this territory, 
and appear to be in a fair way to become established. If this proves to be the case 
the effect of hunting in the early days in the Transvaal and the apparent effect of 
the rinderpest would appear to be quite comprehensible, in spite of the fact that the 
game was not completely eliminated in either case. 

The result of a heavy reduction of game would seem to follow logically, quite apart 
from complete elimination, namely irregular meals at long intervals—perhaps some 
individual starvation—in any case a greatly reduced birth rate. If the birth rate 
were checked at all suddenly, especially during the season of concentration when 
the larvae are deposited within a limited area, the fly’s parasites might be expected 
to destroy a much higher percentage than before. The writer is very much in agree- 
ment with Duke’s contention as to the probable effect of a sudden or “‘ cataclysmic” 
change in the distribution and numbers of game animals. The local distribution of 
G. morsitans in relation to the habits of game animals, even during the time of year 
when the fly is dispersed, is frequently very marked. In this way it appears that the 
effect of shooting and of the rinderpest might be explained, but it is only com- 
prehensible on the supposition, which the writer believes to be justified, that the fly’s 
ability to perceive its hosts is strictly limited, that it 1s incapable of following up a 
trail for any great distance by scent and of attaching itself to and keeping in touch 
day after day with such game as may be present in the infested area. In the latter 
part of the dry season its limitations in locating game, except such as may visit its 
haunts, seem to be obvious, and if, as is probable, this is one of the most important 
breeding periods, the effect of both shooting and rinderpest seems clear enough. 

It may seem improbable that a species of insect should be so ill-provided for taking 
care of itself under varying conditions as the views outlined in this paper suggest, 
but if there is one thing more striking than another about the tsetse it is just this 
disability to adapt itself to a change in its environment, otherwise it would not have 
retreated before the advance of civilisation, in the absence of any direct effort to 
drive it back. 


Summary. 

In the foregoing pages the following tentative views have been put forward in the 
hope that they may receive either confirmation or criticism from other investigators : 

(1). The latter part of the dry season is probably, in general, the most favourable 
portion of the year to the fly. 

(2). The later the onset of the rains the greater the capacity for increase. 

(3). Seasons of unusually heavy rainfall are inimical, at least in certain areas. 

(4). A series of years of low rainfall is favourable. 

(5). Regions where the fly commonly attains the greatest concentration of numbers 
are the regions where it is most liable to be reduced or to die out, namely, parts of the 
infested areas where first-grade foci predominate. 


132 RUPERT W. JACK. 


(6). In Southern Rhodesia the passage of grass fires has, as a rule, no appreciable 
efiect on the numbers of the fly. 

(7). Mopani belts are probably of considerable value to the fly in providing limited 
areas attractive to game in the wet season, where the grass remains short; they 
possibly also afford a convenient refuge from grass fires in the dry season. 

(8). The fly does not migrate under the stimulus of hunger or in company with 
game. 

(9). The fly as a whole does not follow game about infested areas. 

(10). The maximum distance at which a hungry fly readily detects its hosts is a 
short one, possibly less than 100 yards. 

(11). The maximum following distance of the females remains to be determined ; 
the writer is not yet convinced that the females seek animals and human beings only 
for the purpose of feeding, though this appears probable. 


133 


NOTES ON THE MOSQUITOS OF MADAGASCAR, 
MAURITIUS AND REUNION. 
By F. W. Epwarps, 
(Published by permission of the Trustees of the British Museum.) 


Through the kindness of Mons. E. Séguy, of the Muséum d’Histoire Naturelle, 
Paris, I have recently been enabled to examine a considerable number of mosquitos 
from Madagascar and the neighbouring French islands, including the types or co- 
types of all the species described from Madagascar by Ventrillon. As the examination 
of this material has disclosed the existence of several species hitherto unrecorded 
from these islands, as well as some new synonymy, it has been thought desirable 
to record the results at once. In the following pages all the species of mosquitos 
hitherto known from this region are dealt with. The total number is only 28, as 
compared with about 180 in the African fauna, so that it is highly probable that 
further collecting would produce many others. 


Anopheles mauritianus, Daruchy de Grandpré & d’Emmerez de Charmoy. 


Ambatofanghera and Ambohipomana (Bouet, 1905); Tananarive (Nevret and Ven- 
trillon, 1904 ; Bouet 1905). ; 


The commonest Anopheles in Mauritius, according to d’Emmerez. 


Anopheles maculipalpis, Giles. 
Occurs rarely in Mauritius (d’Emmerez de Charmoy). 


Anopheles squamosus, Theobald. 
Cellia tananariviensis, Ventrillon. 

According to Ventrillon this is by far the commonest Anopheles in Madagascar, 
and his statement is certainly borne out by its abundance in the collections which 
Ihave examined. No constant difference was observable between Madagascan and 
African specimens. 


Anopheles pharoensis, Theobald. 


A single specimen, much damaged but undoubtedly this species, from Tananarive 
(Ventrillon, 1905). 


Anopheles funestus, Giles. 
Four females from Moratsiazo, Lac Itasy (Bowet, 1904). 


Anopheles marshalli, Theo. 


Females from Tananarive (Nevret, 1904, and Ventrillon, 1905) and Ambohipomana 
(Bouet, 1905). 


Anopheles transvaalensis, Carter. 
Two females from Tananarive (Bouet, 1905). 


134 F. W. EDWARDS. 
( 


Anopheles costalis (Loew) Theobald. 
Females from Mauritius, where, according to d’Emmerez, the species is common, 
are in the Paris Museum. 


Aedes (Stegomyia) albopicta, Skuse. 

Stegomyra lambert, Ventrillon, Bull. Mus. Paris, x, p. 552 (1904) and Arch. Parasit. 
ix, p. 441 (1905). 

Stegomyva scutellaris, Theobald et auct. (nec Walker). 


Ventrillon’s types included both sexes, so that the identification of S. lamberti 
is beyond doubt, notwithstanding the fact that Ventrillon states that the female 
claws are toothed, whereas in all specimens I have examined they are simple. Hither 
Ventrillon made a mistake, or the species is variable in this respect, which is not 
unlikely. 

This species also occurs at Réunion as shown by specimens in the Paris Museum, 
collected at St. Denis, v-vi, 1913 (Surcouf). It has been recorded from Mauritius, 
where it is‘said to be very common (d’Emmerez de Charmoy). It is surprising that 
it does not seem to occur on the African mainland, since it has such a wide range 
throughout the Oriental region. 


Aedes (Stegomyia) argenteus, Poiret (fasciata, Fb.). 
Culex insatiabilis, Bigot, Ann. Soc. Ent. France, (3) vu, p. 118 (1859). 


Bigot’s description and figurecan apparently only apply to this species, which is 
common on the islands, especially near the coast (d’Hmmerez de Charmoy). 


Aedes (Skusea) cartroni, Ventrillon. 

Stegomyia cartroni, Ventrillon, Bull. Mus. Paris, xii, p. 143 (1906). 

This species, which I had not previously seen, evidently belongs to the sub-genus 
Skusea, although the male is unfortunately unknown. It is very close to S. pembaensts, 
the type of the subgenus, so close indeed that it may eventually prove to be a variety 
only. However in all the examples of S. pembaensis received at the British Museum 
the abdomen is unbanded dorsally, and the scutellar scales are black, while in the 
few specimens of S. cartroni sent the abdominal segments have narrow pale basal 
bands, and the few remaining scutellar scales are white. 


Aedes (Ochlerotatus) nigeriensis, Theobald. 

Culex fowlert, d’Emmerez de Charmoy, Ann. Trop. Med. u, p. 258 (1908). 

Mr. H. F. Carter informs me that he has examined a male of C. fowleri from 
Mauritius, and established its identity with O. nigeriensts, which was already sug- 
gested by the author’s allusion to the two pale spots on the sixth abdominal tergite 
of the female. 

A female in the Paris Museum from Madagascar may be either this species or 
O. fryer, Theo. 


NOTES ON THE MOSQUITOS OF MADAGASCAR. 135 


Orthopodomyia arboricollis, d’ Emmerez de Charmoy. 

Qulex arboricollis, d’Emmerez de Charmoy, Ann. Trop. Med. i1, p. 257 (1908). 

Newsieadina arboricollis, Theobald, Ann. Trop. Med. u, p. 297 (1908). 

This species, which is so far only known from Mauritius, is a typical Orthopodomyra. 
I have examined the types in the Liverpool School of Tropical Medicine and find 
that the female has the very short fourth tarsal joint on the front and middle legs, 
characteristic of this genus, while the male hypopygium is very similar to that of 
the other species. 


Taeniorhynchus (Mansonioides) uniformis, Theobald. 

Females of this species in quite recognisable condition from Ambatofanghera 
(Bouet, 1905). Other specimens, probably of T. (M.) uniformis, were from Morous- 
lava (Dr. Petit, 1901). 


Taeniorhynchus (Coquillettidia) flavus, Ventrillon. 

Culex flavus, Ventrillon (nec Motschulsky), Bull. Mus. Paris, x, p. 550 (1904). 

Culex grandidiert, Blanchard, Les Moustiques, p. 627 (1905). 

Several females, including cotypes of Ventrillon, from Tananarive (Dr. Ventrillon, 
1905). 

This is very close to the African 7’. fuscopennatus, of which it may be a mere local 
form ; the hypopygia are very similar, as is shown by a drawing taken from Ventrillon’s 
male cotype which M. Sécuy kindly sent me; this indicates the clasper as having 
a rather different shape. J’. flavus, however, in all the examples I have seen, is 
readily distinguishable by the coloration of the scales on the wing, particularly the 
apical half. The costa is yellow, with a line of black scales on the outer margin on 
its basal two-thirds ; subcostal and first longitudinal veins predominantly yellow, 
apically entirely so ; stems of fork-cells and tips of veins round wing-tip yellow ; 
remaining veins and fringe mainly black. This description is considerably at 
variance with that of Ventrillon, but an examination of the cotypes showed that 
his was inaccurate. 

T. fuscopennatus differs in having the dark and light wing-scales fairly evenly 
mixed, and also in having the integument of the thorax blackish or dark brown, 
that of 7. flavus being light brown. 

T. aureus, Edw., has an almost greater resemblance even than 7’. fuscopennatus, 
but it again has a rather differently shaped clasper, and has the costa entirely yellow. 

I have adopted the name flavus rather than grandidiert, because the Culex 
flavus of Motschulsky is probably an Aéles, Ventrillon’s later flavus being therefore 
valid in the genus Taeniorhynchus. 


Culex ventrilloni, sp. n. 

2. Head scales narrow, blackish ; a rim round the eyes and a longitudinal band 
down the middle pale golden. Proboscis with a distinct yellow median ring, broader 
below than above. Palpi one-fourth as long as the proboscis, black, with some 
white scales about the middle. Thorax blackish brown, bristles dark. Prothoracic 
lobes with a few narrow golden scales ; mesonotum with golden and dark brown 
scales, the former occurring in an indefinite line in the middle and in some large 


136 F. W. EDWARDS. 


rather irregular patches on each side on the front half. Scutellum pale, with narrow 
pale golden scales. Pleurae with four or five small patches of pale ochreous flat 
scales. Abdomen dark brown, tergites with white basal lateral patches and ochreous 
basal bands, which are broadest in the middle; sternites with dark apical bands. 
Legs black ; all the femora black to the base above, ochreous beneath on the basal 
half. Femora and tibiae all with conspicuous whitish tips, no scattered pale scales. 
Tarsal joints conspicuously ringed with ochreous at the base only ; the rings on the 
last two joits very narrow. Claws simple; empodia large, nearly as long as the 
claws. Wings with brown scales, those in the lateral series linear. First fork-cell 
more than twice as long as its stem, its base nearer the base of the wing than that 
of the second. Cross-veins separated by rather more than the length of the posterior. 

Length, 6 mm. 

Mapacascar : Tananarive, 1904, 19 (Dr. Neiret) and 1905, 292 (Dr. Ventrillon). 

Type in the Paris Museum ; paratype in the British Museum. 

This species is noteworthy for the ornamentation of the head, the basally ringed 
tarsal joints, and the large empodia. It has no near ally among the described African 
and Oriental species, and in the absence of the male it is impossible to say in what. 
subgenus it should be placed. Possibly it may be a Lutzva. 


Culex annulitarsis, Macquart. 

The brief description of Macquart will not apply to any known species. It was. 
referred by Theobald to the synonymy of Stegomyia fasciata, but Macquart’s descrip- 
tion of the legs (hind tibiae with a broad white ring before the tip ; hind metatarsi 
white, with a narrow black ring) makes the reason for this far from obvious. 


Culex (Lutzia) tigripes, Grandpré & Charmoy. 
Tamatave (Jaillet). Described from Mauritius. 


Culex (Culex) quasigelidus, Theobald. 

Pseudoheptaphlebomyia madagascariensis, Ventrillon. 

This synonymy, previously given by me, can be confirmed from an examination 
of the types, although Ventrillon’s description does not agree. Additional specimens. 
are from Ambohipomana (Bouet, 1905) and Tananarive (Waéerlot, 1916). 


Culex (Culex) giganteus, Ventrillon. 

Culex giganteus, Ventrillon, Bull. Mus. Paris, xn, p. 100 (1906). 

Culex nevreti, Ventrillon, Bull. Mus. Paris, xi, p. 103 (1906). 

The original description of C. giganteus and C. neireti are very similar, except as 
regards the size given and the markings of the thorax. Study of the types shows 
that the insects are actually the same, the differences in the description being due to 
inaccurate observation. In particular, Ventrillon’s measurement of 10 mm. 
body length for C. giganteus is a considerable exaggeration, the actual length being 
only about 6 mm., excluding the proboscis. Additional specimens are from 
Ambatofanghera and Ambohipomana (Bouet, 1905). 

The species is a close ally of the African C. annulioris, Theo., differing chiefly in 
the greater extension of the yellow markings of the abdomen; and in some small 
details of the structure of the male hypopygium, notably the shorter and stouter 


NOTES ON THE MOSQUITOS OF MADAGASCAR. 137 


basal arms of the tenth sternites. Perhaps the single specimen of C. annulioris 
recorded by d’Emmerez de Charmoy (Ann. Trop. Med. u, p. 260) may have been 
C. giganteus. 


Culex (Culex) sitiens, Wied. 
Culex ronaldi, d’Emmerez de Charmoy, Ann. Trop. Med. ii, p. 259 (1908). 
Recorded from Mauritius, but not from Madagascar, where however it is very 
likely to occur, since it has a wide distribution from East Africa to Australia. 


Culex (Culex) univittatus, Theobald. 

Heptaphlebomyia montforti, Ventrillon, Arch. Parasit. ix, p. 448 (1905). 

My previous statement as to the identity of C. uwnivittatus and H. montforti is 
confirmed by an examination of the male hypopygium of one of Ventrillon’s 
specimens. 


Culex (Culex) pipiens, L. 

Tananarive (Nevwret, 1904 ; Ventrillon, 1905). 

This record forms a further notable extension of the known range of this species, 
which until recently was thought to be confined to the temperate regions of the 
northern hemisphere. It is now known also from South and East Africa and from 
the Argentine. 


Culex (Culex) ? laurenti, Newstead. 

Some female specimens from Tananarive (Neiret, 1904) are perhaps this species, 
though it is possible that they may belong to the variety of C. pipiens with an 
unbanded abdomen, which is known from East Africa and from the Mediterranean 
region. 

Culex (Culex) fatigans, Wied. 

Culex cartroni, Ventrillon, Bull. Mus. Paris, xi, p. 429 (1905). 

Culex anxifer, Bigot, Ann. Soc. Ent. France, (3) vil, p. 117 (1859). 

The hypopygium of Ventrillon’s type male was mounted and proved it to be 
this species. Other specimens from Réunion (Surcouf). The commonest mosquito 
in Mauritius (d’Emmerez de Charmoy). 

Bigot’s description of C. anxifer is unrecognisable, and he had no type, but 
Blanchard has referred his name to the synonymy of C. fatigans, and there seems 
no sufficient reason to dispute this. 


Culex (Culex) argenteopunctatus, Ventrillon. 

Heptaphlebomyva argenteopunctata, Ventrillon, Arch. Parasit. ix, p. 446 (1905). 

Heptaphlebomyva kingi, Theobald, Ann. Trop. Med. vii, p. 601 (1913). 

A number of females, all in poor condition, some barely recognisable, from 
Tananarive (Ventrillon, Bouet, Guérin Méneville, Waterlot). 

A male in the British Museum named by Ventrillon lacks the abdomen, but 
fortunately there is another, in perfect condition, from the Gold Coast (Dr. A. 
Ingram). I have mounted the hypopygium of this and find that it is a true Culex 
bearing a rather considerable resemblance to C. decens, Theo. The two silvery spots 
on the thorax are remarkable and absolutely diagnostic, being comparable only 
with certain species of Aédes, e.g., punctothoracis, Theo. 


138 F. W. EDWARDS. 


Uranotaenia neireti, sp. n. 

¢S Head scales and integument blackish, a spot of pale blue in the middle in 
front, and another on each side at the eye-margin. Proboscis slightly longer than 
the abdomen, slightly swollen at the tip. Thorax brown, lower half of pleurae 
somewhat lighter. Prothoracic lobes with light blue scales; proepimera bare, 
lower half blackish. Mesonotum with narrow dark brown scales, those on the 
scutellum flat ; no lateral line of flat scales. A spot of pale blue scales in middle of 
pleurae, and a few more scattered on sternopleura ; integument darker on each 
side of the blue spot. Abdomen dark brown above, lighter brown beneath. 
Legs dark brown ; coxae and bases of femora ochreous ; tip of third and whole of 
fourth tarsal joint of hind legs whitish; fifth darker. Tarsi and claws normal. 
Wings with brown scales only, those towards the apex rather narrowly spatulate. 
Upper fork-cell more than half as long as its stem. 

Length, 3 mm. 

MapacascaR: 1 $1 9 (Dr. Ventrellon), without exact locality, labelled “ Ficalbia 
neirett. Type: Ventr.” Type g in the Paris Museum. The female and a second 
male from Tananarive (Ventrillon, 1905) have the blue scales replaced by greyish 
white, probably owing to fading. 


Eretmopodites quinquevittatus, Theobald. 
Eretmopodites condei, Ventrillon, Arch. Parasit. ix, p. 144 (1905). 


No males were present, and the few females in Ventrillon’s collection were rather 
rubbed, but the above synonymy, previously suggested by me, is highly probable. 


139 


NOTES ON THE NOMENCLATURE OF CERTAIN AFRICAN TABANIDAE 
(SUB-FAMILY PANGONIINAB), WITH DESCRIPTIONS OF A NEW 
GENUS AND NEW SPECIES. 
By Major E. E. Austen, D.S.O. 


Some six years ago, it was correctly pointed out by Bréthes (Bull. Soc. Ent. de 
France, 1914, p. 59) that the generic designation Diatomineura, Rond. (Archiv. 
Zool. Anat. Fisiol. ii, p. 84, 1864), is a synonym of Osca, Walk. (Ins. Saund., 
“Diptera, Part i, p. 10, 1850—nec Walk., 1864, nec Stal, 1871). The first species 
mentioned by both Walker and Rondani is Pangonia depressa, Macq. (=Erephopsis 
lata (Tabanus latus), Guér.), and this species is designated by Bréthes as the genotype. 
Erephopsis lata, Guér., of Kertész’s Catalogus Dipterorum (iii, p. 165, 1908), must 
therefore be known henceforth as Osca lata, Guér., and the South African Tabanus 
barbatus, L. (Pangonia barbata, Auct.), and Pangonia fulvifascia, Walk., must be 
transferred to the genus Osca. 

It is not, however, simply a question of substituting Osca for Diatomineura, 
since Osca lata, Guér., is certainly not congeneric with any, or at any rate with the 
majority of the species at present grouped under Diatomineura, sub-genus Corizoneura. 
So far as regards Ethiopian and Oriental species, the difficulty can be met by, as is 
hereby proposed, raising Corizoneura, Rond. (loc. cit., p. 85) to generic rank, desig- 
nating as its genotype the first species mentioned by Rondani under Corizoneura, viz., 
Tabanus aethiopicus, Thunb. (syn. Pangonia appendiculata, Macq.), and erecting 
a new genus, which may be termed Buplex,* for certain Ethiopian species included 
by Kertész under the Corizoneura division of Diatomineura, but not congeneric 
with C. aethiopica, Thunb. The arrangement indicated may be expressed in tabular 
form as follows. 

Eyes bare ; first posterior cell open. 

Ocelli absent ; face, especially in 9, markedly produced (forming a snout-like 
prolongation), with a shining callus, or at least a more or less shining area, on each 
side ; proboscis long or very long, usually much longer than thorax including 
scutellum, and generally horizontal or nearly so; distal extremities of first and 
second joints of front tarsus in g generally each produced above into a tongue- or 
lappet-like process, which, in case at least of second joint, is often of considerable 
relative length be ‘ .. Corizoneura, Rond. 

(Genotype, Tabanus ens Thunb. —syn. a ests aethvopica, Thunb. ; 
Pangoma appendiculata, Macq.). 

Ocelli usually present ; face not produced, merely convex, and without shining 
callus or area on each side; proboscis of only moderate length, shorter than or 
about as long as thorax including scutellum, and generally slanting downwards 
at an angle of 45°; distal extremities of tarsal joints never produced .. as 
Buplezx, gen. nov. 
(Genotype Pangoma suavis, Lw.). 


* Bou-rdné 6, aN Ox-goad. 


140 E. E. AUSTEN, 


Among other species referable to the new genus thus briefly characterised are 
Pangonia brunnipennis, Lw., P. subfascia, Walk., Corizoneura albifacies, Ric., 
CO. dissimilis, Ric., and the new species described below. 

The geaus Corizoneura, Rond., as defined and restricted above, includes, in 
addition to the genotype and the new species described in this paper :—Diatomineura 
virgata, Austen ; D. inornata, Austen ; D. neaver, Austen ; Pangonia sagittaria, Surc.; 
Corizoneura distincta, Ric. ; Pangonia lateralis, Wied. (Fabr. ?); Diatomineura 
hastata, Austen; D. lineatithorax, Austen; D. penetrabilis, Austen ; Corizoneura 
pallidipennis, Ric. ; and C. umbratipennis, Ric. The foregoing all belong to the 
Ethiopian Region, and congeneric with them are the Oriental Pangonia taprobanes, 
Walk. (syn. P. rufa, Macq.), and P. longirostris, Hardw. 

Another change which may as well be made now, since in any case it is almost 
certain to be effected sooner or later, is the replacement of Macquart’s generic 
designation Cadicera by Phara, Walk. (Ins. Saund., Diptera, Part i, p. 9, 1850). 
The latter name, accompanied by a brief diagnosis, was applied by its author to one of 
“* several groups or subgenera” of Pangoma, Latr. ; eighteen species were mentioned 
under Phara, the first three of these being, in the order given, Pangonia melanopyga, 
Wied., P. chrysostigma, Wied., and P. crassvpalpis, Macq. The present writer hereby 
proposes to regard Pangonia melanopyga, Wied., as the genotype of Phara, Walk. ; 
to raise the latter to generic rank ; and to restrict it so as to include besides the 
genotype, among the species mentioned under Phara by Walker, only Pangonia 
chrysostigma, Wied., and P. crassipalpis, Macq., since these three species are 
obviously congeneric. Phara, Walk., therefore, as thus restricted, replaces and 
must be substituted for Cadicera, Macq. (Mém. Soc. Imp. des Sc. Lille, 1854, (2) 
p. 42, 1855), founded for C. rubramarginata, Macq. (loc. cat, p. 23). 


Except where otherwise stated, the types of the new species described in the 
following pages are in the British Museum (Natural History). 


Genus Buplex, nov. 


Buplex fuscinervis, sp. n. 

0,.—Length (2 specimens) 12 to 13 mm. ; width of head 4 to 4:2 mm. ; width of 
front at vertex 0°8 mm. ; length of proboscis 3°4 to 3°5 mm. ; length of wing 12°4 to 
12°5 mm. 

Body, except certain areas at sides of abdomen, black above, with scutellum, lateral 
borders and three narrow longitudinal stripes on scutum, and deep posterior borders to 
abdominal segments smoke-grey* or pale smoke-grey pollinose ; lateral extremities of 
tergites of first (visible) and second abdominal segments smoke-grey, more or less tinged 
with ochraceous tawny owing to the subjacent ground-colour ; wings with base, costal 
and sub-costal cells and stigmatic area ochreous, veins after first longitudinal in distal half 
(from base of discal cell onwards) dark brown, and for most part more or less strongly 
suffused with mummy brown. 

*For names and illustrations of colours used for descriptive purposes in the present 


paper, see Ridgway, ‘‘ Color Standards and Color Nomenclature” (Washington, D.C. 
Published by the Author, 1912). 


NOTES ON THE NOMENCLATURE OF CERTAIN AFRICAN TABANIDAE. 141 


Head : ocelli present ; face, jowls and lower half of front light buff pollinose, clothed 
with pale yellowish hair, occiput smoke-grey pollinose, and likewise clothed with 
pale yellowish hair; upper part of front with a pair of narrow, blackish stripes, 
commencing at ocellar tubercle, descending somewhat lower than middle and 
diverging below, each stripe clothed with short, curving black hairs, interspersed 
with some glistening yellowish hairs ; palpi clothed with yellowish hai, longer on 
proximal than on terminal joint, latter russet or ochraceous tawny, proximal 
joint mouse-grey, russet towards distal extremity ; first and second joints of antennae 
clothed partly with yellowish hair, partly with black hair, first jomt mouse-grey or 
deep neutral grey, second joint russet (third joint missing in case of type). Thorax: 
smoke-grey longitudinal stripes on dorsum extending from front to hind margin of 
scutum, confluent posteriorly ; transverse suture smoke-grey, forming a connection 
between lateral border and outer smoke-grey stripe on each side ; pleurae and pectus 
light greyish olive pollinose ; thorax clothed with yellowish hair, black stripes on 
dorsum clothed posteriorly with fine black hair. Abdomen: black area on first 
(visible) and second tergites restricted to a transverse band occupying middle third 
of anterior half, with rounded posterior angles in each case, and, especially on second 
segment, indented in middle line behind ; lateral margins of fourth and following 
tergites, and lateral extremities of hind border of third tergite ochraceous tawny, 
though normally concealed by hair and smoke-grey pollen ; seventh tergite entirely 
grey ; lateral extremities and hind borders of all tergites clothed with glistening 
hair, longer and ochreous on lateral extremities of second and two or three following 
segments, shorter and paler elsewhere ; black area on second tergite clothed with 
yellowish hair, corresponding areas on two following tergites clothed with short, 
erect black hair; venter smoke-grey, clothed with appressed, glistening, cream- 
coloured hair ; second sternite with a transversely elongate, somewhat reniform, 
fuscous black blotch in middle line, resting on or close to anterior border and confined 
to anterior half of segment, third and three following sternites each with a sooty 
black, transverse band on front border, widely separated in each case from lateral 
margins, and from twice to four times deeper on third than on either of the other 
sternites, these bands clothed with short, erect, black hair. Wangs with auxiliary 
and first longitudinal veins, second longitudinal from base to a little beyond origin 
of third longitudinal, praefurcal portion of fourth longitudinal, and extreme bases 
of fifth and sixth longitudinal veins ochreous ; veins otherwise more or less dark 
brown. Squamae cream-bufi. Halteres maize-yellow, knobs sometimes darker 
(mummy-brown) at base. Legs: coxae and femora deep mouse-grey, clothed with 
yellowish hair ; tibiae and tarsi clove-brown, clothed with minute black hairs, tarsal 
joints sometimes paler towards base. 

South Africa (Cape Province) ; type and para-type from Bizana, East Pondoland, 
1912(—Goodall : : presented by Mr. T. B. Goodall). 

The species described above shows a decided resemblance to Buplex (Pangonia) 
suavis, Lw., but, apart from its smaller size, is distinguishable, inter alia, by the 
presence of the two blackish stripes on the front ; by the outer two of the three 
longitudinal smoke-grey stripes on the disk of the scutum being narrower ; by the 
absence of a brown blotch near the tip of the wing, above the fork of the third longi- 
tudinal vein; and by the veins in the region of the discal cell being suffused with 
mummy-brown. 


142 E. E. AUSTEN. 


Genus Corizoneura, Rond. 
Corizoneura formosa, sp. n. 

o.—Length (4 specimens) 20 to 21:4, mm. ; width of head 6 to 6°4 mm. ; distance 
from upper margin of occiput to anterior extremity of face 5 to 5°25 mm. ; length of 
proboscis 13 to 15 mm. ; length of wing 17-4 to 18 mm. ; wing-expanse 21°5 to 23 mm. 

Dorsum of thorax dark olwwaceous black, with broad smoke-grey lateral borders clothed 
with cream-coloured hair, and with two narrow, smoke-grey, admedian, longitudinal 
stripes ; dorsum of abdomen ochraceous tawny, with distal extremity (fifth to seventh 
segments) infuscated (iron-grey to olwaceous black), a large black median blotch on 
each of the first four visible segments, and fourth segment bordered posteriorly with 
appressed, silvery white and ochreous hair ; venter cinnamon-buff or connamon-coloured, 
with last three segments sometimes mouse-grey and lateral extremities of first visible 
scute neutral grey ; wings with a sepraceous tinge ; processess at tups of firstand second 
joints of front tarsus in 3 very long. 

Head : face bluntly conical, moderately produced ; front and face (except sides of 
facial prominence below, which are clove-brown and shining) pinkish buff pollinose, 
front and borders of face adjacent to eyes clothed with longish, cream-buff hair ; 
occiput smoke-grey, clothed above with cream-buff, below with whitish hair ; jowls 
and basi-occipital region pale smoke-grey, clothed with long whitish hair ; palpi russet, 
dark brown on outer side, proximal joint with a bunch of whitish hair at base below; 
first and second joints of antennae isabella-coloured pollinose, clothed above and 
below with longish hair, generally black or blackish but sometimes mainly cream- 
coloured on first joint, third joint ferruginous or vinaceous rufous, clove-brown 
at tip. Thorax: dorsum, including scutellum, clothed for most part with somewhat 
appressed, cream or cream-buff-coloured hair, often but little visible when regarded 
from above, hair above lateral borders between bases of wings often largely or mainly 
dark brown, pleurae and pectus clothed with fairly long, whitish or yellowish white 
hair. Abdomen: except on first (visible) tergite, on which median blotch extends’ 
to or is but narrowly separated from hind margin, black median blotches, which 
rest on front margin, do not reach hind border ; posterior angles of first and second 
tergites clothed with yellowish or whitish hair, hind border of second tergite with 
a patch of glistening, appressed, ivory-yellow or cream-bufi-coloured hair in middle 
line, light-haired hind border of fourth tergite expanded in middle line and at each 
extremity, lateral extremities of sixth and seventh tergites generally clothed with 
glistening silvery white hair, lateral extremities of second to fourth tergites inclusive, 
except as already stated, clothed with black hair, ochraceous tawny area of first and 
second tergites clothed mainly with minute, glistening, appressed, ochreous hairs, 
corresponding area on third and fourth tergites clothed with minute black or blackish 
hairs, fifth and sixth tergites (except lateral extremities in case of latter) clothed 
with black hair; venter clothed with minute, appressed, glistening, cream-bufi- 
coloured hairs, fifth sternite sometimes largely clothed with minute, black or blackish 
hairs. Wings: veins sepia-coloured, adjacent membrane suffused with same colour 
at base, and to a less extent at level of proximal extremity of discal cell. Sqwamae 
waxen ochraceous buff or pale orange-yellow, fringed with minute yellowish hairs. 

Halteres light chestnut-brown, tips of knobs light buff or light ochraceous buff. 
Legs: coxae neutral grey, clothed with whitish hair, which is longer on front pair ; 


NOTES ON THE NOMENCLATURE OF CERTAIN AFRICAN TABANIDAE. 143 


femora russet (extreme tips ochraceous buff), clothed mainly with whitish or yellowish 
white hair, antero-inferior area of front pair with minute, erect, blackish hairs ; front 
and middle tibiae ochraceous buff, clothed with minute, glistening, yellowish or 
ochreous hairs, hind tibiae and hind tarsi russet-brown, clothed with black hair, 
distal extremities of hind tarsal joimts blackish ; front tarsi ochraceous buff or 
ochraceous tawny, distal extremities of last three joints dark brown, body of second 
joint very short, tongue-like process at its distal extremity very long (longer than 
following joint), process at end of first joint overlapping and closely applied to 
first two-thirds of following joint ; first joint of middle tarsi cream-buff, remaining 
joints dark brown or mummy-brown, blackish brown at tips. 

South Africa (Cape Province): type and three para-types from Deelfontein, 
7.xi1.1902 (presented by Colonel Sloggett, A.M.S.—now Lneut.-General Sir A. T. 
Sloggett, K.C.B., K.C.M.G.). 

This handsome species presents a decided resemblance to Corizoneura aethiopica, 
Thunb., another inhabitant of the South African portion of the Ethiopian Region, 
but can at once be distinguished, inter alia, by the presence of the conspicuous, 
smoke-grey, admedian, longitudinal stripes on the dorsum of the thorax. 


Corizoneura schwetzi, sp. n. 

3 9.—Length, ¢ (6 specimens) 16°4 to 18 mm., 2 (6 specimens) 16 to 18°6 mm. ; 
width of head, g just over 5 to 5°4 mm., 9 52 to 5°75 mm.; distance from upper 
margin of occiput to anterior extremity of face, f 4:2 to 45 mm., 9 45 to 48 
mm. ; width of front of 2 at vertex 0°6 to 0°75 mm. ; length of proboscis, 3 15 to 
15°25 mm., 9 11°75 to 13°75 mm. ; length of wing, ¢ 15 to 15°2 mm., 2 15°5 to 
17°25 mm. 

Dusky species, with base of abdomen paler; dorsum of thorax dark brownish olive, 
with a broad, indistinct, blackish brown longitudinal stripe along each side above lateral 
margin, a narrow, elongate, pale spot (composed of Naples yellow or cream-buff hair) 
above base of each wing wm front of postalar callus, and a short but conspicuous stripe 
of black hawr between base of costa and humeral callus on each side ; dorsum of abdomen 
with first two (visible) segments connamon-buff or tawny olive in 3, ochraceous tawny 
m 2, 3 with a blackish brown median triangular spot at base of second segment, tergites 
of third and following abdominal segments blackish brown or black, fourth segment 
posteriorly with a conspicuous transverse band of appressed, glistening, silvery white 
hair ; venter abruptly bicoloured, proximal portion as far as base of third (visible) 
stermite, wory-yellow, cream or cream-buff-coloured, remainder blackish brown ; wings 
strongly and uniformly tinged with sepia. 

Head drab pollinose, posterior orbits paler (light greyish olive above, smoke- 
grey or pale smoke-grey below), shining callus on each side of face below antenna 
black, sharply defined, sides of face below calli more or less dark brown or blackish 
brown owing to ground-colour showing through pollinose covering, lower border 
of sides of face in ¢ shining blackish brown, deeper posteriorly, distal extremity 
of face in @ shining blackish brown, sparsely clothed with drab pollen, in both 
sexes a more or less distinct, somewhat triangular, olive-buff or pale olive-buff, 
pollinose spot on each side of lower part of front, between base of antenna, margin 


of eye and shining callus ; front in Q with following series of dark brown marks— 
(687) D 


144 E. E. AUSTEN. 


a transversely oblong spot occupying ocellar region of vertex but not extending to 
eyes, a pair of somewhat guttate spots in centre of front, indistinctly connected 
with foregoing spot but likewise not in contact with eyes, and an oblique spot on 
each side below, extending from base vf antenna to eye, above the light (olive-buff 
or pale olive-buff) spot already mentioned ; front in ¢ clothed with black hair, 
in 2 with shorter blackish hair, mingled with paler hair in upper portion, posterior 
orbits in both sexes fringed above with yellowish hair and below with longer whitish 
hair, basi-occipital region and jowls densely clothed with fairly long, whitish hair, 
lower borders of sides of face clothed posteriorly with black or blackish hair, more 
conspicuous in g than in 2; palpi russet-brown in J, russet in 9, terminal segment 
(elongate and very narrow in 4) clothed with minute black or blackish hairs, proximal 
segment, especially m Q, clothed below with longer hair, brownish or blackish 
brown anteriorly, yellowish or whitish posteriorly ; antennae russet, first and second 
joints more or less smoke-grey pollinose, clothed with black hair. Thorax: dorsum 
clothed anteriorly with silky, tawny olive or ochreous hair, posteriorly, including 
base of scutellum, with fine, erect, black hair, postalar calli, and scutellum, except 
as stated, clothed with Naples yellow or cream-buff hair, hairy covering of central 
portion of dorsum visible only when viewed from side ; pleurae and pectus clothed 
with longish hair, for most part cream-buff (whitish on propleurae) in colour, a tuft 
of black hair (less conspicuous in Q, in which sex it is often much reduced) below 
base of wing on each side ; sternopleurae in 3 sometimes clothed mainly with black 
or blackish hair. Abdomen: hind border (distal third or rather less) of second 
(visible) tergite clothed with minute, appressed hairs, which are silvery white or 
whitish at and towards lateral extremities of segment, and sparser and more yellow- 
ish (often glistening cream-buff) in vicinity of median line (owing to paler ground 
colour, whitish transverse band thus formed is, except at lateral extremities, less 
‘conspicuous than corresponding band on fourth segment); base of first (visible) 
tergite with a blackish brown, median area, extending bevond scutellum, but not 
or barely reaching hind margin except in middle line in 4, in which sex it is more 
or less distinctly connected with triangular spot on second segment; blackish 
brown median triangle on second tergite in ¢ with its base resting on or close to 
anterior margin, and its apex reaching or extending somewhat beyond middle of 
segment ; third tergite sometimes irregularly paler (cinnamon-brown) at base and 
on hind border ; hairs in silvery white transverse band on fourth tergite somewhat 
yellowish in vicinity of middle line; seventh tergite in Q, as well as frequently 
hind borders of fifth and sixth tergites, often dull fuscous ; lateral extremities, 
or at least posterior angles, of fifth and sixth tergites each clothed with a prominent 
tuft of silvery white hair; posterior angles, as well as in 9 hind margin of first 
(visible) tergite clothed with shining ochreous or pale ochraceous orange hair ; 
dorsum of abdomen except as already stated clothed with minute, appressed, black 
hairs ; first (visible) sternite bare, second sternite clothed with minute, appressed, 
glistening cream-coloured hairs (anterior border of second sternite fringed with fine, 
erect, black or blackish hair, shorter and less developed in @ than in 3, in which 
sex fine, recumbent, black hairs are also present among the cream-coloured hair, 
especially towards lateral extremities of the scute), blackish brown portion of venter 
clothed with black hair, among which a few pale hairs are occasionally present 


NOTES ON THE NOMENCLATURE OF CERTAIN AFRICAN TABANIDAE. 145 


on one or more of fourth and following segments. Wangs: appendix to anterior 
branch of third longitudinal vein, though of variable length, as a rule relatively 
somewhat long. Squamae isabella-coloured. Halteres mummy-brown, knobs 
in 2 often paler (cream-buff) at tip. Legs: front coxae neutral grey, clothed with 
cream-coloured or cream-buff hair, middle and hind coxae deep neutral grey or 
dark neutral grey, clothed with black hair, hind coxae in 9 also with some yellowish 
hairs ; hind femora in ¢ and bases of front and middle femora in same sex blackish 
brown, front and middle femora in ¢ except at base russet-brown or cinnamon- 
brown, all femora in ¢ clothed with black hair, femora in Q paler, and clothed 
largely with ochraceous buff or ochreous hair as well as with black hair ; coloration 
of tibiae and tarsi and of their hairy covering alike in both sexes, front and middle 
tibiae ochraceous buff or ochraceous tawny, clothed with minute, appressed glisten- 
ing ochraceous buff hair, hind tibiae and hind tarsi blackish brown, clothed with 
black hair, tips of second and two following tarsal joints clothed below with 
ferruginous hair, similarly coloured hair also largely present, at least in 9, on under 
side of first tarsal joint, front and middle tarsi ochraceous tawny, clothed above 
with black hair, last joint in each case, as well as distal extremities of preceding 
joints mummy-brown, processes at tips of first and second joints of front tarsi in 
S$ of moderate size, in neither case reaching distal extremity of following joint. 

Belgian Congo (North Katanga) and Tanganyika Territory. Typical series 
from Kakanu (between 15 and 16 miles south of Kisengwa, R. Lomami), N. Katanga, 
vi. 1918 (Dr. J. Schwetz). Type of 3, type of 9, 36 gd and 2 9 para-types, in Musée 
Royal d’Histoire Naturelle de Belgique (Brussels) ; 6 J and 6 2 para-types, in 
British Museum (Natural History)—presented by M. G. Severin ; 1 9, from Tangan- 
yika Territory (formerly German East Africa), 30° 55’ E. Long., 2° 5’ S. Lat., 
16. vi. 1916 (Dr. G. D. H. Carpenter), in British Museum (Natural History), presented 
by Imperial Bureau of Entomology. 

This fine species, with which the author has much pleasure in connecting the 
name of its discoverer (the well-known student of tsetse-fly bionomics in North 
Katanga), was met with by the investigator in question in large numbers in the 
vicinity of Kakanu.* On 6th June 1918, in a belt of forest several hundred metres 
in breadth and about 33 miles from Kakanu, Dr. Schwetz’s native carriers succeeded 
in catching some 2,000 specimens of C. schwetzi, from 80 to 90 per cent: of which 
were males.f Unlike Corizoneura inornata, Austen (see below), which is found 
in the open, C. schwetzi does not occur outside the forest. According to Dr. Schwetz,t 
in the case of the present and the following species (C. inornata, Austen) at least, the 
labium itself is the piercing organ, and in the act of biting is thrust by the insect 
deeply into the skin of its victim. It is therefore interesting and possibly suggestive 
to note that, in the Q taken by Dr. Carpenter in Tanganyika Territory, the labium 


* Of. Schwetz, ‘‘ Dix Jours d’Observations sur les Moeurs de la ‘ Pangonia zonata’ 
et de la ‘ Pangonia oldii’ (Deuxiéme Note)”: Revue Zoologique Africaine, vii, pp. 92-106 
(1919).—Cf. also the earlier paper by the same author, ‘* Quelques Observations Pré- 
liminaires sur les Moeurs de la ‘ Pangonia zonata’’’: ibid., pp. 46-54. In both of the 
memoirs cited Corizoneura schwetzi is referred to as Pangonia oldii, while the species 
termed Pangonia zonata is really Corizoneura inornata, Austen. 


+ Cf. Schwetz, loc. cit., p. 103. 
t See below, p. 147. 
(687) D2 


146 EK. E. AUSTEN. 


only projects 7°5 mm. beyond the extremity of the clypeus, exceeding the length 
of the labrum-epipharynx and the other mouth-parts by little more than the labella, 
while the proximal portion of the labium is bent backwards underneath the head, 
and beneath the cleft between head and thorax forms an angle of 45° with the distal 
portion. 


Although superficially presenting a decided resemblance to Pangonia oldi, Austen, 
Corizoneura schwetz can be distinguished from that species inter ala by the processes 
(entirely wanting in P. oldw) at the tips of the first and second joints of the front 
tarsi in the 9; by the much greater development of the dark spot (often scarcely 
more than vestigial in P. oldi) at the base of the second abdominal tergite in the 
- same sex ; and in both sexes by the short stripe of black hair on each side of the 
dorsum of the thorax, in front of the base of the wing. 

From Corizoneura inornata, Austen, apart from obvious differences in coloration, 
especially the sharply bicoloured venter of the species just described, C. schwetzi is 
distinguished by the inferior development of the processes at the tips of the first and 
second joints of the front tarsi in the g. Whereas in C. inornata 3 each of these 
processes is so long as to project beyond the tip (excluding the process in the case 
of the second segment) of the following joint, in C. schwetzi 3 neither process reaches 
the tip of the succeeding joint. 


Corizoneura inornata, Austen. 
Diatomineura mornata, Austen, Bull. Ent. Res. 1, p. 282 (1911). 
This species was described from a single 9, obtained in September 1907, in 


S. Katanga, Belgian Congo, between Bunkeya and Kambove, at an altitude of 
3,500 ft., by Dr. Sheffield Neave. The kindness of Dr. J. Schwetz, in presenting 
to the British Museum (Natural History) a large number of specimens of both sexes, 


now renders it possible to indicate the distinctive characters of the ¢. 


C. inornata, Austen, g.—Apart from ordinary secondary sexual characters, 
agreeing generally with the 2 except as follows. Head: hair clothing jowls and 
basi-occipital region often hoary or nearly white; first two joints of antennae 
clothed mainly with black hair. Thorax: pleurae on each side with a tuft of black 
hair below base of wing. Abdomen: first (visible) tergite with a median brownish 
black area at base, projecting somewhat beyond scutellum, but not reaching hind 
margin ; second tergite with a conspicuous, median, brownish black, triangular 
spot resting on front margin, and varying in size in different individuals, but not 
extending beyond middle of segment, if so far; fifth and sixth tergites mainly 
brownish black, blackish brown or clove-brown, each of the two preceding tergites 
often with an ill-defined median blotch of same colour occupying anterior two-thirds. 
Legs : front and middle femora clothed largely with black hair, at least towards base, 
hind legs clothed mainly with black hair, processes at tips of first and second joints. 
of front tarsi very long, in each case projecting beyond end of following joint 
(excluding process in case of second segment). 

In the papers already referred to,* under the name “ Pangonia zonata,” Dr. 
Schwetz has furnished a series of interesting field notes on the behaviour of this 


* P, 146, note.* 


NOTES ON THE NOMENCLATURE OF CERTAIN AFRICAN TABANIDAE. 147 


species, which, at the end of May 1914 and at the close of the same month and 
beginning of June 1918, was found by him in great abundance in the vicinity of 
Kakanu, N. Katanga, at and about the flowers of Acanthus montanus (termed by 
the natives ‘‘ Nafimbia’’), which grows in sheets in open spaces outside the forest. 
According to Dr. Schwetz, the existence of C. inornata is “ intimately connected ” 
with that of the flowers in question, the nectar of which is imbibed by both sexes, 
though the females also suck blood on occasion. It is interesting to note that, 
having in a number of cases observed the process of sucking blood by females of this 
and the foregoing species (Corizoneura schwetzi, Austen), Dr. Schwetz states that, 
in these PANGONIINAE at any rate, the actual piercing organ is the proboscis (labium) 
itself, which is “‘ evidently sufficiently rigid ”’ to penetrate the human integument, 
and is driven by the insect “ fairly and squarely into the skin to the extent of 
one-third or one-half of its length. . . .” * 


Genus Pangonia, Latr. 
Pangonia discors, sp. n. 


©.—Length (1 specimen) 19°6 mm. ; width of head 5°6 mm. ; width of front at 
vertex 0°75 mm.; distance from upper margin of occiput to anterior extremity 
of face 45 mm. ; length of proboscis 5°5 mm. ; length of wing 17°6 mm. 

Deep black, somewhat shining ; first (visible) and second abdominal tergites densely 
covered with pale gull-grey pollen, and thickly clothed with closely appressed, silvery 
white hair ; area beneath scutellum free from pollen and clothed with black hairs, some 
black hairs also present in middle line on anterior border of second tergite, latter likewise 
exhibiting a broad, median, triangular area (its base resting on front margin, its apex 
directed backwards and reaching beyond middle of segment) which, like front border of 
same segment, has a blackish look owing to pollen on it being thinner than elsewhere ; 
lateral extremities of second tergite clothed with black hair ; wings mummy-brown, 
proximal half (as far as base of discal cell) and stigma pale orange-yellow or light orange- 
yellow. 


Head : ocelli wanting ; face moderately prominent, front above relatively some- 
what narrow ; area from middle of front to anterior margin of clypeus olive-buff, 
upper half of front fuscous black, sides of face clove-brown, occiput and basi-occipital 
region smoke-grey or pale smoke-grey, jowls clothed with yellowish cream-coloured 
hair ; palpi clove-brown, terminal joint elongate, tapering to a point and somewhat 
curved ; first and second joints of antennae dark mummy-brown, clothed with 
minute black hairs mixed with some minute yellowish hairs, first joint short, expanded 
portion of terminal joint cinnamon-brown (terminal portion missing in case of type). 
Thorax: postalar calli chestnut-brown ; dorsum including scutellum clothed with 
short black hair, which on front border of scutellum is interspersed with minute, 
glistening, appressed, golden hairs, a few golden hairs also on upper portion of 
swelling occupying depression at each end of transverse suture, hair on pleurae and 
pectus entirely black or fuscous black. Abdomen: third (visible) and following 
tergites clothed with appressed, black hair ; venter, except second sternite, clothed 
with appressed, black or brownish black hair, ventral surface of second segment | 


* Of. Schwetz, Rev. Zool. Africaine, vii, pp. 101-102 (1919). 


148 E. E. AUSTEN. 


clothed with minute, appressed, glistening, yellowish white hairs, fore border and 
lateral extremities clothed with black hairs. Wangs: transition from orange- 
yellow proximal to mummy-brown distal portion sharply marked, at least in case 
of type. Squamae ivory-yellow. Halteres ochraceous buff, stalks and knobs 
towards base brownish. Legs: coxae and femora dark brown or blackish brown, 
clothed with black hair, which on under side of hind femora is mixed with a certain 
number of minute, glistening tawny hairs, and on anterior surface of front coxae 
with a few golden hairs ; tibiae and tarsi russet, clothed for most part with minute, 
appressed, glistening, ochraceous tawny hairs, last three joints of tarsi dark brown 
above. 

Angola (J. J. Montevro). 

Pangona discors resembles and is closely allied to the East African P. beckeri, 
Bezzi, but is readily distinguishable oyving to, inter alia, its more prominent face and 
narrower upper part of the front, the entire absence of white hair on pleurae, postalar 
calli and front coxae, and the distal portion of the abdomen being entirely black and 
covered with black hair, instead of having the tip ochraceous tawny and clothed 
with glistening ochraceous orange or ochreous hair. 

Pangonia lautissima, sp. n. 

¢g 2.—Length, g (9 specimens) 17 to 19 mm., Q (3 specimens) 16 to 17°4 mm. ; 
width of head, g just under 5 to 5'4 mm., 9 5 to 5°6 mm.; width of front of 9 at 
vertex just under 1 to 1-2 mm. ; length of proboscis, g just under 4 to 4°25 mm., 
2 3°4 to 4-2 mm. ; length of wing, 9 14 to 15:2 mm., 9 14 to 15°6 mm. 

Shining black ; basi-occimital region clothed and lower halves of posterior orbits 
Jringed with orange-buff hair, and patches of similarly coloured hair on pleurae ; 
wings with base and a deep anterior border extending to end of third costal cell ochraceous 
orange, and remainder of surface uniformly brownish black, with a strong purplish 
metallic sheen. 

Head black, frontal triangle in § and region of subcallus (area immediately above 
bases of antennae) in Q shimmering silvery white pollinose, a similar pollinose — 
patch (clothed with a few whitish hairs, and usually more distinct and sharply 
defined in 9 than in Q) on each side of upper part of face in both sexes ; face in both 
sexes tumid below antennal prominence, then indented or somewhat receding, not 
produced into a nose-like prolongation, front in Q deeply furrowed ; occiput pallid 
neutral grey pollinose, clothed with whitish hair, posterior orbits silvery white, 
their upper halves fringed behind with minute blackish hairs ; palp: and antennae 
black, proximal joints of both sparsely clothed with short, black or blackish hairs, 
third joint of antennae from certain aspects appearing dark olive-grey, mouse-grey 
or brownish grey pollinose. Thorax: dorsum including scutellum clothed with 
short black hair, humeral calli inconspicuously neutral grey pollinose and clothed on 
sides in front with pale orange-buff hair, postalar calli fringed posteriorly below with 
orange-buff hair; pleurae on each side with a thick tuft of orange-buff hair below 
humeral callus, and more posteriorly with two further tufts of similar hair arranged 
somewhat in the shape of a wide V, of which the anterior branch fringes the hind 
margin of the mesopleura, while the posterior runs back to the squamae ; pleurae 
except as stated, and pectus except in front of front coxae clothed with black hair, 
pectus in front of front coxae neutral grey pollinose, clothed with pale orange-buff 


_ NOTES ON THE NOMENCLATURE OF CERTAIN AFRICAN TABANIDAE, 149 


hair. Abdomen: first six (visible) tergites in both sexes each with its lateral fourth on 
each side (rather more in case of 2) clothed with close-set, appressed, minute, glistening 
smoke grey hairs (extreme lateral extremities of seventh tergite in 9 clothed with 
hairs of same kind), so that dorsum of abdomen exhibits on each side a broad grey 
stripe, which in certain aspects contrasts fairly sharply with remainder of surface, 
which is clothed with minute appressed black hairs and thus forms a broad, longi- 
tudinal, median, black stripe ; first (visible) tergite clothed with greyish pollen 
on each side at base, second tergite in both sexes on each side with a roughly semi- 
circular whitish pollinose spot on hind margin, from most points of view concealed 
by the smoke-grey hair, but clearly visible when abdomen is looked at obliquely 
from behind, in @ a pair of similar but smaller whitish pollinose spots on hind 
margin of each of the two following tergites also; venter clothed with minute, 
appressed, glistening, smoke-grey hair, last two sternites, and sometimes median 
area of hind border of preceding sternite also, clothed mainly or entirely with black 
hair. Wings: extreme base of costa and first longitudinal vein brownish black ; 
ochraceous orange base extending into bases of basal and anal cells, similarly 
coloured anterior border including upper margin of first basal and proximal fourth 
of first submarginal cell, thence tapering obliquely to end of third costal cell ; 
veins within ochraceous orange area similarly coloured, elsewhere dark brown. 
Squamae light orange yellow. Halteres: stalks and knobs mummy-brown above and 
below, tips of knobs cream-buff. Legs black and clothed with black hair, front 
coxae anteriorly dark neutral grey pollinose, thinly clothed towards base with longish 
pale orange-buff hair. 

Tanganyika Territory (formerly German East Africa) : Itigi, iv, 1917 (Dr. G. D. H. 
Carpenter). Type of 3, type of 9, and 2 9 para-types, taken 18.iv.1917; 7 g 
para-types, taken 15.iv.1917, “on composite flower”; 1 3 para-type, taken 
6.iv.1917, “ among low herbage.” All foregoing presented by Imperial Bureau of 
Entomology, which retains possession of six additional para-types, taken by Dr. 
Carpenter at same time and place as specimens already enumerated. 

The extremely striking and unusual-looking Tabanid just described belongs to 
the group of the genus Pangonia that includes P. elongata, Ric., P. becker, Bezzi. 
and P. discors, Austen, but is readily distinguishable from all three species by, 
apart from its sharply defined wing-markings and other characters, its unbanded 
abdomen and the patches of orange-buff hair on the pleurae. The coloration of 
the wings, in conjunction with the shining black body, would seem to suggest that 
P. lautissima possibly mimics some species of wasp (perhaps Rhynchium cyanopterum, 
Sauss.), and it is worthy of note that the same colours, though differing widely 
in extent and arrangement in the case of the wings, are exhibited by “ Pangonia” 
mesembrinordes, Surc., of which the type was also obtained in Tanganyika Territory 
(Amani). The latter species, however, was incorrectly assigned by its describer to the 
genus Pangonia, and really belongs to a new genus allied to Thriambeutes, Griinb. 


Pangonia carpenteri, sp. n. 

Q.—Length (8 specimens) 15 to 16°6 mm.; width of head 4°25 to just under 
5 mm. ; width of front at vertex 0°6 mm. ; distance from upper margin of occiput 
to anterior extremity of face 3°5 to 4 mm. ; length of proboscis 11°75 to 13°75 mm. ; 
length of wing 13°5 to 15 mm. 


150 E. E. AUSTEN. 


In Q sex, at any rate, a somewhat sombre-coloured species looking more like a small 
or medium-sized Corizoneura than a Pangonia, and not unlike a smaller and more 
dusky form of Corizoneura hastata, Austen, of Portuguese E. Africa.—Face moderately 
produced ; dorsum of thorax olivaceous black, with a pair of broad, paler (greyish olive), 
narrowly separated or sometimes confluent, admedian, longitudinal stripes ; dorsum of 
abdomen shining blackish brown, on each side of base rather more than lateral third of 
first (visible) tergite ochraceous tawny, a similar area on each side of second tergite, 
or of second and third tergites, usually russet or chestnut-brown, lateral borders, posterior 
angles, and (at least in part) hind borders of second and fourth tergites clothed with 
glistening silvery white hair; wings strongly tinged with sepia; legs for most part 
ochraceous tawny. 

Head: ocelli wanting ; face and front clothed with yellowish grey or isabella- 
coloured pollen, and front thinly covered with yellowish hair, lower half of front, 
above antennal prominence, with an ill-defined, shining black, median longitudinal 
mark, upper half of front usually streaked with mummy-brown or dark brown along 
each side, while a narrow mummy-brown streak, which starts from base of antenna 
on each side, runs obliquely upwards and outwards, and joins corresponding eye 
just above base of antennal prominence ; below antennae, a shining black transverse 
band unites and includes the shining black facial calli; jowls and basi-occipital 
region clothed with long, whitish hair ; occiput greyish olive or smoke-grey pollinose, 
sparsely clothed with cream-coloured hair, which also forms a short fringe behind 
posterior orbits above, lower portion of posterior orbits fringed with whitish hair ; 
palpi elongate, proximal joint russet-brown or deep mouse-grey, sparsely clothed 
with whitish hair on outer side and below, terminal joint russet, somewhat expanded 
in middle, clothed with minute black hairs ; first and second joints of antennae 
pinkish cinnamon or cinnamon-buff pollinose, both sometimes clothed with black 
hair though in some specimens hair on first joint is for most part yellowish, second 
joint usually with outstanding black hair above and below, third joint orange- 
cinnamon. Thorax: admedian stripes on dorsum entire, extending from front 
margin to prescutellar groove, outer border of each stripe paler in front of transverse 
suture ; dorsum including scutellum thickly clothed with fine yellowish hair, with 
which in some specimens on central part of posterior portion of scutum a few fine 
black or blackish hairs are intermixed ; postalar calli and lateral margins of dorsum 
behind transverse suture fringed with whitish hair, pleurae and pectus thickly 
clothed with similar hair. Abdomen: central portion of base of first (visible) 
tergite olivaceous black, basal angles of same tergite more or less distinctly neutral 
grey or dark neutral grey ; rather less than median third of second tergite and a 
broad median triangle on each of following tergites, in each case with its base resting 
on hind margin and its apex in contact with front margin, and on fourth and following 
tergites with its base expanded laterally so as to include entire hind border of segment, 
dull dark olive-grey ; olive-grey median area on second tergite (sometimes that on 
third tergite also) indented on each side ; second tergite at base with a median, 
semicircular, dead black spot, or in some cases with a pair of smaller spots, narrowly 
separated in middle line by a dark olive-grey longitudinal stripe ; first (visible) 
tergite with a median patch of glistening yellowish or pale yellowish hair, and 
clothed elsewhere with ochraceous hair; second tergite, except on black spo 


NOTES ON THE NOMENCLATURE OF CERTAIN AFRICAN TABANIDAE. ld1 


(or spots) on which hair is usually black or ochreous, clothed with minute, appressed, 
glistening silvery white hairs, which however, at least on each side of median olive- 
grey area, are often largely replaced by similar ochreous hairs ; lateral borders 
of fourth and following tergites, as well as (at least in part) hind border of fourth 
tergite, clothed with glistening, silvery white hair, similar hair also usually visible 
on lateral margins of third tergite, towards posterior angles ; dorsum except as 
stated clothed with appressed black hair ; venter isabella-coloured or light brownish 
olive, clothed with minute, appressed, whitish or yellowish white hairs. Wangs: 
veins mummy-brown ; first posterior cell variable as usual as regards distance 
from hind margin at which it 1s closed, sometimes closed on margin itself, or even 
in one or other wing narrowly open. Squamae cream-buff. Halteres: knobs 
fuscous, stalks cinnamon-buff. Legs: coxae neutral grey, clothed with whitish 
hair, anterior and inferior surfaces of femora clothed, at least in part, with black 
hair, femora elsewhere clothed with yellowish hair, tibiae and tarsi clothed with 
minute, appressed, ochreous hairs, extensor surfaces of hind tibiae and hind tarsi 
clothed, at least in part, with black hair ; front femora blackish brown at base and 
sometimes also on greater part of under side, joints of front tarsi often mummy- 
brown or dark brown at tips above, those of hind tarsi similarly marked, or some- 
times entire upper surface of hind tarsi, except base of first joint, dark brown. 

Tanganyika Territory: Itigi, iv, 1917 (Dr. G. D. H. Carpenter). Type and 
3 para-types, taken 18.1v.1917; 1 para-type, taken 15, iv, 1917, “ on low herbage ”’; 
1 para-type, taken 6.iv.1917, “ hovering while feeding from composite flower ; 
looking much lke a Bombylius.” All foregoing presented by Imperial Bureau of 
Entomology, in whose possession are two other para-types, taken by Dr. Carpenter 
at same time and place as specimens already mentioned. 

So far as itis possible to judge from the 9 alone,this species, which is named in honour 
of its discoverer and does not resemble any African Pangonia hitherto described, 
presents, apart from the venation, all the characteristics of a Corizoneura, to which 
genus it would have been assigned were it not that its posterior cell seems normally 
to be closed before reaching the wing-margin. Should the 3 prove to have processes 
at the tips of the first and second joints of the front tarsi, Pangonia carpentert, despite 
the transitional character of its venation, would more fittingly be placed under 
Corizoneura, so long as the independence of the latter be maintained. 


Genus Thriambeutes, Griinb. 
Thriambeutes fuscus, sp. n. 

§.—Length (1 specimen) 11:14 mm.; width of head 4°25 mm.; length of wing 
10°5 mm. 

Dorsum of thorax sepia-coloured, with traces of a faintly marked, paler, longitudinal 
median stripe im front of transverse suture, pleurae and pectus mummy-brown ; abdomen 
uniformly blackish brown ; wings mummy-brown, with a clear oblique transverse streak, 
commencing on anterior transverse vein (its base extending from commencement of 
lower border of distal fourth of first basal cell to proximal extremity of first posterior cell), 
wmeluding rather more than proximal third of discal cell, proximal extremity of fourth 
postervor cell, distal extrematy of second basal cell, and upper border of proximal two- 
thirds of fifth posterior cell, but not reaching hind margin ; legs blackish brown or black, 
middle tarsi cream-buff, last joint and tips of preceding joints cinnamon-brown. 


152 E. E. AUSTEN. 


Head black or blackish brown, occiput dark neutral grey pollinose, basi-occipital 
region thinly clothed with fine blackish brown hair ; ocelli present, enlarged facets 
of eyes very coarse, area occupied thereby same as in ¢ of genotype (Thriambeutes 
singularis, Griinb., of Togoland and 8S. Nigeria), 7.e., small facets confined to a deep 
lower border and a narrower hind border of uniform width running up to ocelli ; 
palpr blackish brown and clothed with fine hair of same colour, terminal joint 
elongate and curved but not conspicuously swollen, considerably smaller and 
narrower than in g of genotype; antennal protuberance large and prominent, 
considerably larger and more prominent than in 3 of genotype; first joint of 
antennae blackish brown, short, swollen, cylindrical, and clothed like second joint 
with blackish brown hair, second and third joints sepia-coloured, expanded portion 
of third joint rather broad. Thorax and abdomen thinly clothed with fine blackish 
brown hair. Wdngs: anal angle and lower region of distal extremity short of actual 
tip paler than elsewhere with exception of clear, transverse streak, a close scrutiny, 
when wing is viewed against a light back-ground, revealing beyond clear streak 
an ill-defined transverse band, which appears somewhat darker than remainder of 
surface ; stagma well developed, elongate, cinnamon-brown when seen against a. 
light background. Sguamae blackish brown. Halteres: knobs ivory yellow, stalks 
sepla-coloured. Legs: coxae, femora and tibiae clothed with blackish brown or 
blackish hair, middle as well as front tibiae swollen (front and hind tarsi, and hind 
tibiae missing in case of type). 

South Africa, Bechuanaland Protectorate : N’Gami Country, 1897 (Sir Frederick 
Lugard, G.C.M.G., C.B., D.S.O.). 

The species characterised above is readily distinguishable by its wing-markings. 
alone, apart from all other characters, from Thriambeutes singularis, Griinb. (the only 
other member of its genus as yet described), in which moreover the body as well as. 
the head and its appendages are in the § uniformly tawny. 

So far as it is possible to judge from a photograph, which is all that 1s at present 
available for comparison, what appears to be another ¢ of Thriambeutes fuscus 1s 
in the possession of Mr. R. W. Jack, Government Entomologist, Southern Rhodesia, 
and was taken by him in November 1914, in Sebungwe District, Southern Rhodesia, 
on the jacket of a companion. In Mr. Jack’s specimen, however, the clear streak 
in the wing reaches the hind margin, while the margin of the anal angle, and a further 
portion of the hind border embracing part of the distal extremity of the second 
submarginal cell and the distal extremities of the first three posterior cells are also. 
hyaline. 


153 
APHIDIDAE OF PERSIA. 
By F. V. THEOBALD. 


Aphis buxtoni, sp. nov. (fig. 1). 

Alate viwiparous female. 

Head and thorax dark ; abdomen pale, apparently pale green, with dark lateral 
spots. Antennae, cauda and cornicles dark. Legs with mid and hind femora dark, 
front pair paler ; apices of tibiae and the tarsi dark. Antennae shorter than body, 
rather thick ; basal segment larger and paler than second ; third much longer than 
fourth and about as long as the sixth, with 18-20 small round pale sensoria over its 
whole length ; fourth segment very slightly longer than fifth, with 5-8 sensoria ; 
fifth with 4-7 sensoria of unequal size, exclusive of the usual subapical one ; sixth 
with the rather thick basal area less than one-third the length of flagellum, in some 
however almost half its length ; all the segments from the third imbricated. yes 


Fig. 1. Aphis buxtoni,sp.n.: A, B, alate 9 ; 
a,, lateral bladder-like papilla; b,, lateral 
papilla; C,D, apterous ¢; d,, lateral papilla. 


large. Proboscis reaching to the third coxae. A large pale bladder-like papilla 
on each side of pronotum and five pairs of pale rounded papillae on sides of abdomen. 
Cornicles about as long and as thick as third antennal segment, cylindrical, markedly 
imbricated, not reaching to the level of the cauda. Cauda bluntly pointed, about 
half the length of the cornicles, spinose ; with numerous fine hairs, curved apically. 
Anal plate spinose, with fine pale hairs. Tibiae with numerous fine, pale, short 
hairs. Wings with normal venation; veins and stigma brown, the veins very 
faintly and narrowly clouded on each side. Length, 2-2°2 mm. 
Apterous viviparous female. 


Pallid, with dark head, cornicles, cauda and anal plate, also dark antennae and 
mid and hind legs, the fore pair paler. Antennae shorter than body, rather thick ; 
basal segment larger than second, both very dark ; third paler at base, longer than 
fourth and about same length as sixth ; fourth a little longer than fifth, the latter 
with the usual sub-apical sensorium ; sixth with basal area nearly half the length of 


154 F. V. THEOBALD. 


flagellum ; segments 3-6 imbricated. Eyes dark, of moderate size. Proboscis 
reaching to the second coxae. Cornicles cylindrical, longer than third antennal 
segment, imbricated. Cauda blunt and rather short, not half the length of the 
cornicles, spinose, with numerous pale hairs ; broader than cornicles ; the hairs curved 
apically. Anal plate spinose, with long pale hairs. The rather thick set legs have 
the coxae and trochanters darker than the rest of the legs; a few short hairs on 
the femora, many on the tibiae. Length, 2 to 2°5 mm. 

Food-plant: Umbelliferae. 

Persia : Enzeli, Caspian Coast, 6.v1.1919 (P. A. Buzton). 

Types in the British Museum. 

Described from a single perfect alate female and several apterae. Oneof the latter 
shows the cornicles not reaching the cauda, others passing it. It 1s a very marked 
species, the black cornicles, cauda, etc., showing up prominently against the pale body. 
The antennae are short, as in the genus Cavariella and the cauda is markedly spimose. 
I can see no trace in the mounted apterae of any lateral pronotal or abdominal 
papillae, as seen in the alate female. The larval stages are more uniform in colour, 
but in succeeding instars the blackness of the cornicles, etc., gradually becomes more 
pronounced. 

This species was found in association with an ant that has been identified by Mr. 
W. C. Crawley as Lasius emarginatus, Latr., var. nigro-emarginatus, Forel. 


Myzus mespiliella, sp. nov. (fig. 2). 

Apterous viviparous female. 

Green or yellowish green ; antennae of same colour as body, dusky at apex of 
fifth segment, the sixth dusky, with a more or less darkened area at the junction of 
basal portion and flagellum. Eyes deep reddish-black. Proboscis of same colour 
as body, dark at apex. Legs same colour as body, apex of tibiae and tarsi dusky. 
Cornicles and cauda of same colour as body ; in some the former seem a little darker. 
Anal plate darker than cauda. 


Fig. 2. Myzus mespiliella, sp. n.: A, B, apterous 2; C, D, alate 9. 


Antennae longer than body ; basal segment larger than second ; third segment 
longer than fourth, but not quite so long as the sixth ; fourth a little longer than 
fifth ; the sixth about as long as 4+ 5, with moderately long flagellum ; a few 
short hairs on third and fourth. Proboscis rather long, reaching to or just beyond 
the third coxae. Cornicles slightly longer than fourth antennal segment, cylindrical, 
slightly broadening at the base, markedly imbricated. Cauda reaching not quite 
to the level of the cornicles, less than half their length, narrowly triangulate or 
pointed, spinose, with two pairs of lateral hairs. Anal plate spinose, with several 
long hairs. Tibiae with numerous small hairs, becoming longer near apex. The 


APHIDIDAE OF PERSIA. 155 


vertex shows two short and two longer slightly capitate hairs. A few short blunt 
lateral abdominal processes. Length, 1°5-1°8 mm. 

Alate viviparous female. 

Only imperfect specimens sent. Apparently dark and of reddish hue with dark 
head, thorax, cornicles and cauda ; legs paler, with dark apices to tibiae and dark 
tarsi; two basal segments of antennae dark, third pale. The abdomen has dark 
lateral and dorsal spots and the proboscis is pale towards the tip, but extreme apex 
dark. The basal segment of antennae larger than second ; the third with 6—7 large 
round sensoria in a line. Eyes large and deep reddish-black. The black cornicles 
are cylindrical, shorter, but a little thicker than third antennal segment, slightly 
expanding basally and not quite reaching the level of the cauda, markedly 
imbricated. Cauda black, more than half the length of the cornicles, more or less 
constricted near the middle, spinose, with long, fine hairs; blunt apically. Anal 
plate black, with long pale hairs. Wings large, venation normal. The abdomen 
shows four small dark blunt lateral processes before the cornicles and one between 
the cornicles and cauda. Length, 1°8-2 mm. 

Food-plant : Medlar (Mespilus). 

N. W. Persia: Enzeli, 29.iv.1919 (P. A. Buzton). 

Types in the British Museum. 

Described from several perfect apterous females, but both alatae damaged, 
neither showing complete antennae. ‘The sensoria on the third segment are marked, 
The only other Mespilus species is Van der Goot’s Myzus mespili, which is very 
. distinct. The vertex and lobes of the apterae and the basal segment of the antennae 
approach to some extent those of the genus Phorodon, but in the alate stage it is a 
distinct Myzus. It was found on the young twigs of the medlars, not doing much 
harm. Ants were found in attendance. 


Aphis punicae, Passerini (fig. 3). 
A. punicae, Pass., Aphid. Italicae, pp. 32 et 42, n. 26 (1863). 
Alate viviparous female. 


Antennae shorter than body ; two basal segments dark and a dark area at apex 
of fifth and around the sensoria on sixth, or all dark except base of third segment : 


Fig. 3._ Aphis punicae, Pass.: A, B, C, alate 9; 

D, E, F, apterous 2; dj, dark areas on antennae. 
basal segment wider but no longer than second ; third nearly as long as the sixth, 
with 6-8 sensoria spread over its whole length, two frequently smaller than the 
others ; fourth segment longer than fifth, the latter with the usual subapical 
sensorium ; the sixth about as long as 4 +5; fourth to sixth imbricated. Cornicles 


a 


156 F. V. THEOBALD. 


dark, about as long as fourth antennal segment and thicker, cylindrical, imbricated, 
not reaching to the tip of cauda. Cauda pale, shorter than the cornicles, with three 
pairs of lateral hairs, curved apically and arising from prominent tubercles. Anal 
plate darker than cauda. Jemora of mid and hind legs dark, except just at base ; 
front pair paler ; apices of tibiae and the tarsi dark. Wings normal. Length, 1-15 mm. 

Apterous viviparous female. 

Antennae shorter than body, pallid, except for a small dark area at apex of fifth 
segment and one around sensoria on sixth; basal segment wider but scarcely 
longer than second ; third a little shorter than sixth, but longer than fourth ; fourth 
and fifth about equal; sixth with flagellum about twice as long as basal area. 
Cornicles about as long as third antennal segment, broadening basally, pale, dark at 
apex, not reaching as far as cauda. Cauda pale, with three hairs on each side, 
curved at their tips and arising from marked tubercles ; not quite as long as cornicles. 
Anal plate dark. Marked pronotal lateral tubercles and one each side of body 
before the hind legs. Apices of tibiae and the tarsi dark. Proboscis not reaching 
to third coxae. Length, 1-15 mm. 

Food-plant : Wild Pomegranate. 

N.W. Persta: Tula Road, Talish District, 8.W. Coast of Caspian, 9. vi. and 
5. vil. 1919 (P. A. Buxton). Ivraty (Passerint and Theobald). 

Apparently the apterae only were described by Passerini. The alatae seem to 
vary in the colour of the antennae in the mounted specimens, some are pale with 
two dark areas, others all dark except the base of the third segment. The specimens 
taken by Mr. Buxton were on the twigs, attended by ants (Cremastogaster scutellaris 
schmidti, Mayr, and Tapinoma erraticum mgerromum, Nyl.). Passerimi records it 
from Punica granatum and P. sylvestris. The Aphis punicellae I described from 
Egypt on P. granatum (Bull. Ent. Res. vi, p. 125) is quite a distinct species. 


Lachnus pyri, Buckton (fig. 4). 

L. pyri, Buckt., Indian Museum Notes, iv, p. 271, pl. xvi (1899). 

Taken on pears at Karmanshah, W. Persia (4. xu. 18) by Mr. P. A. Buxton. 

This species was described by Buckton (with notes by E. E. Green) from pears in 
Ceylon. The Persian specimens agree with the Pear Lachnid I have from Ceylon 
and answer generally to Buckton’s short description. I append some characters of 
the antennae, etc. 


‘ey 


Fig. 4. Lachnus pyri, Buckt.: A, antenna of alate 2; a, basal segments ; 
b, eye; B, cornicle; C, lst to 3rd antennal segments of apterous 9. 


APHIDIDAE OF PERSIA. 157 


Alate viviparous female. 

Antennae of six segments; first two large and dark, third the longest, with 
13-15 round sensoria of unequal size and a few hairs ; fourth about as long as fifth, 
with 3 sensoria ; fifth longer than sixth, with one large subapical sensorium ; sixth 
with ‘nail’ shorter than basal area and a large sensorium at its base, a few 
moderately long hairs. yes large. Cornicles dark, of normal form, hairy. 

Apterous viviparous female. 

Basal segments of antennae large and dark; third segment with 3-4 uniform 
round sensoria situated near apex. Cornicles as in alate 9. A marked median 
abdominal tubercle. 

The specimens sent by Mr. Buxton were all very much damaged. Another 
Lachnid (Nippolachnus piri, Matsumura) occurs on Pirus sinensis in Japan, but 
is very distinct (vide, “ A list of the Aphididae of Japan, with descriptions of New 


Species,” Journal of the College of Agriculture, Tohoku Imperial University, 
vi, pt. 6, p. 382, July 1917). 


e. 
‘ 


a ee ros 
« "A a 


= 


Lae ies 
Tie Ate 
*, CTP A ey 


159 


THE CAMPAIGN AGAINST PHYTALUS SMITHI IN THE 
COLONY OF MAURITIUS. 
By H. A. Tempany, D.Sc., 


Director of Agriculture, 


and D. d’EMMEREz de CHARMOY, 


Entomologist. 
(Pate, V.) 


The following paper contains an account of the campaign which has been in pro- 
gress against Phytalus smithi during the past 8} years. The history of the pest 
in Mauritius and the measures which have been adopted for its control are of interest 
both from an entomological and from an administrative point of view. 

Published accounts of the pest are contained (1) in a monograph, prepared by one 
of us at the time of its first appearance in the Colony, and printed in Mauritius ; 
(2) ina paper on the introduction of Tiphia parallela into Mauritius, from Barbados, 
for the purpose of the control of the insect, published in the Bulletin of Entomologi- 
cal Research (iii, pp. 93-102), also published subsequently as a Bulletin of the Depart- 
ment of Agriculture, in Mauritius ; (3) in the annual administrative reports on the 
Department of Agriculture in Mauritius for the years 1913-1918 ; and (4) in special 
reports on the progress of the work of control which have been prepared from time 
to time for presentation to the Government and to the Board of Agriculture in 
Mauritius. 

As, however, the information is scattered, and many of the reports and papers are 
difficult of access to the general reader, the following summarised account has been 
prepared for publication. . 


Historical. 

The first record of the appearance of Phytalus smithi as a pest in Mauritius occurred 
in the month of July 1911, on lands of Mon Rocher Estate adjoiming the Royal 
Botanical Gardens, Pamplemousses. Once the pest had been detected, it became 
apparent that damage on a very considerable scale was being occasioned thereby, 
many fields of both plant and ratoon canes being found to be almost totally des- 
troyed by the grubs. 

The insect was apparently imported into Mauritius, either from Barbados or South 
America, probably in earth containing rooted sugar-canes. The exact date of its 
introduction is uncertain, but it is probable that it must antedate its discovery as 
a pest in the Pamplemousses district by five or six years at least. 

Its observation in Mauritius was followed by prompt and energetic measures for 
its control, which have been continued up to the present time. The effect of these 


(687) E 


160 H. A. TEMPANY AND D’EMMEREZ DE CHARMOY. 


measures has been to circumscribe to a large extent the infested area, and to mitigate 


the damage caused thereby. Had these measures not been applied, it seems reason- 


ably certain that the cane-growing industry, at any rate in the lower regions of the 


island, would have sustained much more serious damage than has actually been 


experienced. 


At the time of the first outbreak, no properly constituted authority existed for 
dealing with administrative measures directed to the control of plant pests and 
diseases, and consequently two committees were appointed, charged with the super- 


vision of the work of control, one being a consultative and the other an executive 


body. At the same time the present Entomologist of the Department of Agriculture, 


who then held the office of Curator of the Mauritius Institute and Museum, was 
appointed temporarily for executive work in connection with the operations, the 


work being attached, for administrative purposes, to the Department of Forests. 


On the formation of the Department of Agriculture in 1913, the control of these 


and other operations against plant pests and diseases was transferred definitely 
to the control of this Department and has remained thereunder ever since. 


The Habits of the Beetle and the Damage done by it. 
The character of this Melolonthid beetle and the nature of the damage occasioned 


by it are now too well known to need any detailed description, and for a complete 


- account of the various stages in its life-history reference may be made to the report. 
on Phytalus smith by one of us (D. d’Emmerez de Charmoy), previously referred to. 


Only one generation is produced per annum. Though the period of the greatest 


emergence does not last more than 3 to 4 months, from November to January or 


February, these insects can be found in the adult stage all the year round in varying 


numbers, according to the season, the development being to some extent dependent 
upon climatic conditions. Soil temperature and soil moisture are factors influencing 
the duration of the pupal and larval stages, as well as the incubation of the eggs. 


The following table brings out the difference existing between insects reared at: 


Phoenix, at an altitude of 1,311 ft., and those reared at Pamplemousses, where the 
altitude is 208 ft., the temperature being in the latter district much higher than in 


the former. 


Egg stage 

Larva: active Lif 
inactive—prepupal 

Pupa 


prior to emergence from soil .. 
first copulation after emergence 
Imago 4; copulation af 2 
laying perod .. Ae 

after the laying period 


No. of days 


Minimum | Maximum 


5 23 
210 240 
30 45 
18 25 
10 15 
10 15 
60 75 
30 120 
30 90 


Remarks 


Maximum at 


Phoenix in 
September. 
Minimum = at 
Pamplemous- 
ses in Decem- 

ber 


THE CAMPAIGN AGAINST PHYTALUS SMITHI. 161 


It would appear that climate is a most important factor in the control of the 
pest ; to exemplify this it may be stated that the insect was first detected in the year 
1907 at Reduit at an elevation of 1,100 feet, where the climate is distinctly cooler 
than at Pamplemousses. The insect has been kept under close observation at this 
point for a number of years past and has not up to the present shown any signs of 
becoming a serious pest. In this connection mention may be made of the very 
recent discovery of the beetle on a small area at Joli Bois in the Savanne district of 
the island at an elevation of about 900feet. Atthis particular point it seems probable 
that the insect was introduced from Pamplemousses in planting materials trans- 
ported from forest nurseries formerly existing in Pamplemousses Gardens, for the 
purpose of restocking Government forests adjacent to this point. No planting 
material has been sent out from these nurseries since the discovery of the pest at 
Pamplemousses in 1911, but just prior to this, important consignments of such 
materials were delivered at Riviere du Poste Railway Station, which is in the centre 
of the infested zone and undoubtedly is the focus from which the infestation 
originated. The fact that during an interval of eight years the pest has extended 
only to an insignificant extent is further evidence of the effect of climate on its 
incidence. 

The extent of the infested area in the Pamplemousses district now measures 
approximately twelve square miles. When the pest was first detected in 1911, the 
infested area measured only three square miles, the increase in the intervening period 
being due to outward spread. Hnergetic measures, described in a later section of 
this paper, are now being taken with a view to circumscribing its further extension. 
The Royal Botanical Gardens are approximately in the centre of the infested zone 
and are apparently the nucleus from which infestation originally started. 


Remedial Measures. 

The remedial measures that have been applied to combat the pest may be grouped 
under the following heads :— 

(a) the collection and destruction of adult insects on emergence ; 

(b) the digging out and destruction of larvae in badly infected fields ; 

(c) census and survey work designed to ascertain the location and distribution 
of foci of infestation throughout the infested area ; 

(d) the introduction and dissemination of the natural parasitic enemies of the 
pest ; 

(e) legislative measures directed towards preventing the spread of the pest to 
uninfested regions, and affording the necessary powers to the authorities 
to carry out protective and destructive work in infested areas ; 

(f) experimental work on other possible means of control. 

The control work in the Pamplemousses area has assumed very considerable 
proportions, and the local headquarters for dealing with the work have for some 
years been established in the Royal Botanical Gardens, In the following paragraphs 
an account is given of the administrative and practical methods employed under 
each of the above sub-heads, after which information is supplied concerning the 
results of the work. 

(673) E2 


162 F. A. TEMPANY AND D. D EMMEREZ DE CHARMOY. 


Collection of the Insects. 

The collection of insects by hand has proved one of the most effective methods 
of controlling the pest, and in this connection organised work under Government 
supervision on a considerable scale has been carried out systematically each year. 

The methods adopted have comprised the collection of insects by the general public, 
combined with collection by gangs in localities where the infestation is light. In 
regard to public collections the principle observed is the purchase of the collected 
beetles at rates dictated by the abundance of the insects at any particular period, 
as gauged by the number of insects captured ; a sliding scale of payments has been 
worked out in this connection, and the rates paid are fixed in accordance therewith. 
The work of collecting usually commences in the month of October and continues 
uninterruptedly till March. The months of greatest emergence are December and 
January, during which period the daily catches not infrequently run into several 
millions. 

The capture of the beetles commences at about 6.30 p.m. and continues up to 10.30 
p-m. The habit of the insect lends itself to this particular form of control measure, 
inasmuch as at nightiall it emerges from the soil, where it lies buried during the day, 
takes a short flight to the nearest bush or tree, and remains there for hours nibbling 
slowly at the leaf on which it has settled. The imsects are undisturbed by lights 
and are readily collected by searchers who look for them with lanterns. 

Beetles caught each night are received, measured and purchased at Pamplemoussses 
Gardens on the following morning, and immediately after measurement destroyed 
by fire. During busy seasons the work of measuring beetles occupies many hours. 
The method employed in computing the number of captured beetles is by means of 
measurement in containers of standard size and known capacity. To facilitate the work 
of collecting beetles, purchasers are licensed by the Agricultural Department, who buy 
beetles from the general public at certain specified points on the night they are col- 
lected and resell to the cfficer in charge the next morning ; while many estates also 
follow the same practice, in some cases paying slightly enhanced premiums in order 
to attract collectors, and also supplying oil for lanterns, collecting receptacles, &c.- 

With regard to collection of sects by gangs as opposed to public collection, 
this form of control has of necessity to be practised in regions where the infestation 
is light and beetles are not numerous, since under the system of purchase, beetle 
hunters are obviously attracted only to localities in which insects are plentiful, 
while the scale of operations renders impossible any disciimination in the matter 
of prices in relation to the localities in which captures are effected. 

Two quite distinct cases occur in which collection by gangs is indicated; the 
first is that of places well within the infested area at which the beetles are not numer- 
ous ; the second is in the case of the peripheral zone, where infestation is light and 
where slow outward spread to previously uninfested regions is taking place. 

This outward spreading constitutes one of the most serious aspects of the situation. 
By 1917 the increase in the infested area had become so marked that special measures 
to restrict the spread became necessary. With this object in view the periphery 
of the infested zone was resurveyed in 1917; following this the periphery was 
divided between a series of patrols, and to each patrol a special collecting gang was 
assigned. Hach gang consists of from 8-10 labourers under the charge of a head- 
man ; it is provided with powerful acetylene lights to assist in the detection of 


THE CAMPAIGN AGAINST PHYTALUS SMITHI. 163. 


insects, and is required to patrol the section allotted to it on a depth of } a mile at 
least once in each night, and to collect insects found occurring on this area. The 
gangs are regularly inspected by a travelling inspector, when attendances are 
verified, captures of insects checked, and the general working of the gang examined. 
As the result of work on these lines during two seasons there appears reason to believe 
that the rate of outward spread is becoming considerably reduced. 

The collection of beetles during busy seasons affords part time employment to 
large numbers of people, and at such times the actual number engaged may total 
several thousands. The largest number of beetles captured on any one night 
amounts to 4,600,000, which was recorded on 12th January 1919. Emergence of 
beetles usually takes place in flights, large numbers appearing simultaneously for 
a short period and then rapidly diminishing. It is usual for two or more flights 
to be recorded in one season, and on occasions when beetles are numerous it is 
remarkable, after nightfall, to observe the country-side dotted with the lanterns 
of collectors. 

The transport of the beetles from the points of collection gave rise some time ago to 
fears concerning possibilities of reinfestation owing to their escape from unsuitable 
containers. A special form of container, intended to preclude the escape of captured 
insects deposited there, has in consequence been devised and efforts,made to 
induce all persons engaged in the work of beetle collection to provide themselves 
therewith. 

Daily returns of captures of beetles and of larvae are supplied to the Head Office ; 
by means of these, detailed control of the work is maintained, and it is found possible 
in this way to detect readily localities which show signs of being especially heavily 
infested and if requisite special attention can then be devoted to them. 

From the outset the collection and destruction of larvae, by digging them out of 
the soil in badly infested areas, has formed an essential part of the control measures 
adopted. As at present conducted, it is principally combined with survey work 
and is carried out during those months of the year when beetle emergence is at a 
minimum and the larvae are attaining full growth, wz., May to August. With the 
oradual spread of Tiphia parallela, the introduced parasite, it has been found 
advisable to restrict the destruction of larvae in localities in which this parasite has 
established itself, as the indiscriminate destruction of larvae in these circumstances 
may do more harm than good, owing to the risk of destroying those that are para- 
sitised. 

Census work in the infested area has been systematically carried out for some 
years past. Under this method in every field in the zone 10 to 15 trial holes of a 
depth of 8 inches are dug in every acre, the larvae found in each hole are counted, 
and an infestation rate per acre is worked out. The results are subsequently 
tabulated in the form of a plan, a system of tinting being adopted to indicate varying 
degrees of infestation. Following this, orders are issued to estate proprietors to 
dig out the larvae on fields which show more than 20,000 larvae per acre. The 
prescribed fields are subsequently investigated again after the digging operations 
have been completed, and if the work has not been satisfactorily performed, the 
Department of Agriculture is empowered to repeat the process and to charge the 
estate owner with the cost. ; 


164 H. A. TEMPANY AND D. D EMMEREZ DE CHARMOY. 


The Introduction of a Natural Parasitic Enemy of the Pest. 

The identification by Nowell, in 1912, of Tephia parallela as the principal insect 
enemy of Phytalus smithi in Barbados led to a series of attempts being made between 
the years 1913-1915 to introduce the insect into Mauritius. A full account of the 
efforts made in this direction are contained in the paper on this subject previously 
referred to, published in the Bulletin of Entomological Research. 


Since the publication of the paper m question Tiphia parallela has established 
itself firmly at several points in the infested zone and at these points has already 
exercised a very marked influence on the incidence of the pest. The effect was first 
seen in 1917 at Esperance estate, where, as the result of the action of the parasite, 
the total number of adult insects captured in the season 1917-1918-showed a very 
marked reduction over that of the previous year, the reduction being still more 
marked in the year 1918-1919. In the year 1918-1919 similar reductions became 
visible in the case of Maison Blanche estate and also on the group of small properties 
classed under the head “Small Planters’ which occur in this vicinity. During 
the present season, though completed figures are not available, there is evidence 
to show that a similar reduction has become evident at Beau Plan Estate; while 
at Esperance where the insects were formerly present in great numbers they are 
now only met with occasionally. 

The following figures for the number of insects captured at these pots during 
the years 1916-17, 1917-18 and 1918-19 illustrate these remarks :— 


Year Esperance . MaisonBlanche Small Planters 
1916-17 21,419,586 — a 
1917-18 7,650,461 1,895,752 8,322,130 
1918-19 518,320 937,950 4,303,625 


The spread of Tiphia throughout the infested area has been systematically 
assisted by the liberation of insects in places where they did not previously exist. 
An important point, which has materially assisted in the dissemination of the insects, 
is the existence of the wild plant Cordia wnterrupta (Herbe Conde), the sweetish 
exudation from the leaves of which provides food for the Tvphia adults. It is also 
noteworthy that the plant attracts Phytalus ; i consequence planters are now 
growing the Cordia around the borders of the cane-fields, and this has materially 
facilitated both the capture of Phytalus and the dissemination of Tiphia. 


Legislative Measures. 

Legislative measures for the control of plant pests and diseases are provided 
under Ordinance No. 8 of 1911. 

This ordinance provides for the proclamation of any prescribed area as being 
infected with plant disease, including both insect and fungous pests, and for the 
making of general or special orders for the following purposes :— 

(a) for prescribing or regulating the destruction, removal, uprooting, disposal 
or treatment of plants and products of a vegetable nature within an infected 
district, area or place ; 

(b) for prescribing or regulating the cleansing and disinfecting of infected 
places or areas, or parts thereof ; 

(c) for prescribing the period within which it will not..be lawful to plant or 
replant with any plant whatsoever, or with any particular kind of plant, 
the whole or any portion of an infected district, area or place ; 


THE CAMPAIGN AGAINST PHYTALUS SMITHI. 165 


(d) for prohibiting or regulating the movement of persons, animals, earth, 
soil, manure, vegetable product or other things into or out of a district, 
area or place declared to be a place infected or suspected of being infested 
with a plant disease ; 

(e) for the appointment, pay and regulation of the duties of persons to be 
charged with the execution of any of the provisions of or orders made 
by virtue of this Ordinance ; 


(f) generally for the purpose of in any manner preventing the spreading within 
the island of any plant disease. 


It further provides for the removal of such restrictions in the event of infection 
being stamped out, for the entry on occupied lands of duly authorised persons for 
the purpose of examining plants and for carrying out remedial, disinfective or 
prophylactic measures, for the notification of plant diseases and for penalties for 
contravention of the provisions of the Ordinance. 


Under this Ordinance, Proclamation No. 10 of 1913 prescribed the area infested 
with Phytalus smithi, and made certain orders in relation thereto. 


In 1918 the spread of the pest and experience with its control rendered desirable 
the rectification of the prescribed area and slight modifications in the order ; accord- 
ingly the above proclamation was superseded by Proclamation No. 42 of 1918 the 
terms of which are as follows :— 


(a) the removal of living specimens of Phytalus snuthi either larvae, pupae or 
adults from the area prescribed is prohibited. This, however, shall not 
apply to the transport of living insects in captivity by properly authorised 
officers of the Department of Agriculture for purposes of scientific study. 

(b) the removal of all earth, manure, leaf mould, decayed vegetable matter and 
all living plants or parts of plants in soil is prohibited from the area pre- 
scribed, but the above shall not apply to fruits or vegetables growing above 
the ground and detachable from the tree or stem on which they grow. 

(c) cane-tops for propagation purposes in such limited quantities as may be 
decided upon by the Director of Agriculture may, after examination and 
disinfection if necessary at Pamplemousses Gardens or at such convenient 
point as the Director of Agriculture may select, be removed in sealed 
packages from the infested area. 


(d). the Director of Agriculture or other person deputed in writing by him may at 
any time inspect sugar-canes or other crops within the specified area and issue, 
if considered necessary, an order in writing containing instructions as to the 
cutting, reaping, disinfection or other treatment of such sugar-canes or crops 
and as to the disinfection or treatment of the soil in which they are growing. 
Owners, managers or occupiers shall within 48 hours of such service 
commence to carry out the treatments ordered, and failing to carry them 
out, the Director of Agriculture or person aforesaid may cause such treatments 
to be carried out at the expense of the owner, manager or occupier notified. 


(e) the Director of Agriculture may plant, raise or cultivate such trees, shrubs 
or plants as may be deemed expedient for the purpose of attracting or 
trapping, and may spray or otherwise treat the same for the purpose of 
destroying the pest. These trees, shrubs or plants shall be protected by 
the owner, manager or occupier of the land on which they are planted or 
cultivated for such time as the Director of Agriculture may consider 
necessary. 


166 H. A. TEMPANY AND D. D’EMMEREZ DE CHARMOY. 


The measures above indicated have on the whole provided adequate machinery 
for the control of the pest. One point, however, requires mention, namely the risk 
of artificial dissemination of the pest by interested parties, as a result of the policy 
of buying the beetles. To guard against this a supplementary Ordinance was passed 
in 1918, whereby any person wilfully disseminating a plant disease is rendered 
liable to imprisonment without the option of a fine. 


Other Methods of Control. 

In the early days of the outbreak, a considerable number of experiments were 
tried on methods of control, other than those enumerated above. These included 
the trial of various stomach and contact poisons and also the injection of carbon 
bisulphide and other poisons into the soil. The pest, however, exists in such numbers 
and on such very extensive areas that the cost of methods of this description renders 
their employment prohibitive for the most part. The principal methods which 
have been tried are described below. 

Numerous experiments were undertaken at the outset with a view to finding an 
efficient insecticide, moderate in cost and easily applicable, that would permit of 
the control of the pest, more particularly on young plantations, as these suffered most 
from its attacks. 

Powder insecticides, such as Paris green and vaporite, proved unsuitable on 
account of the labour expenses involved in their application. Neither potassium 
cyanide nor carbon bisulphide could be utilised, not only on account of the expense, 
but owing also to their harmful effects on plants. Liquid insecticides, involving 
the use of kerosene, creoline and carbolic acid, though rather expensive, gave more 
satisfactory results when dealing with young plantations and first ratoons, but their 
application could not be generalised, in most cases for want of the water needed in 
their preparation. 

Experiments were also tried with the object of finding some substance that would 
attract the larvae. Good results were obtained with refuse cane-cuttings and tops. 
These were split in halves and buried in small furrows 2 or 3 inches deep. Every 
fortnight the furrows were opened, the cuttings turned over and the larvae removed, 
the same cuttings being replaced and revisited. This method not only allows of 
the destruction of the pest at a comparatively cheap rate, but also prevents the 
larvae from entering the planting holes when in search of food. 

Other substances such as molasses, ashes, scums, &c., have been experimented 
with, but without success ; attempts were in addition made to infect adults and larvae 
with cryptogamic diseases. Those affecting larvae proved contagious, but could 
not be transmitted experimentally ; while the one that attacks the beetle and was 
introduced from Porto Rico (Metarrhizium anisopliae) gave poor results. It was 
found subsequently that this fungus existed already in the Colony, attacking various 
species of SCARABAEIDAE, but did not exert any very great influence in checking 
their spread. 


The Number of Insects captured and the Results of the Campaign. 

The statistics for the number of insects captured afford the best indications of 
the total prevalence of the pest. These are given below in tabular form, the figures 
showing both insects and larvae destroyed on all the properties concerned. 


167 


THE CAMPAIGN AGAINST.PHYTALUS SMITHI. 


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168 H. A, TEMPANY AND D. D EMMEREZ DE CHARMOY. 


The figures for the 1919-20 campaign are as yet incomplete, but the data so far 
accumulated appear to give reason for the expectation that the total captures re- 
corded will be markedly less than those of the three preceding years. It will be 
observed that during the years 1916-17 to 1918-19 the total number of insects 
captured per annum has remained practically stationary ; before that date they 
showed a fairly steady annual increase, mainly due to the increase of the infested 
area ; while it is clear that the observed check to this increase is the the result of the 
control measures which have been applied, the actual result experienced is the 
outcome of a number of superimposed factors. As already poimted out, marked 
reductions have taken place in the intensity of infestation at Esperance, Maison 
Blanche and in the case of small planters, largely as the result of the introduction 
of Tiphia parallela. In localities where Tiphea has not yet established itself, the 
tendency is on the whole still slightly towards an increase in numbers, while in 
addition there is the factor of outward spread, and it is estimated that if Tuphoa 
had not been introduced, the insects captured in 1918-19 would probably have 
totalled 100,000,000. It would appear that collection methods by themselves have 
proved efiective in reducing the incidence of the pest and have greatly mitigated the 
damage done. 

Collection and the work of T7phia parallela combined appear to afford reasonable 
hope that in infested areas they will ultimately render the incidence of the pest 
comparatively unimportant. The problem of preventing the spread of the pest 
to uninfested areas is, however, a difficult one and if this is to be effected, a continua- 
tion of some measure of quarantine and peripheral patrol collection seems essential. 

Even if Tiphia becomes established throughout the whole of the infested area, 
it can never by its nature establish a complete control of Phytalus, since for its 
existence it is dependent on the production of a supply of beetles each year. It is 
moreover powerless to check completely the spread of the beetle, since Tiphia will 
naturally congregate where Phytalus is plentiful, that is to say, well within the 
infested area, while the pest spreads at the circumference where the attraction for 
Tiphia is least. Whether in the event of the successful establishment of Tiphia 
throughout the infested area such a slow spread can be regarded in the end with 
equanimity, it is impossible at the present stage to say. It is, however, quite clear 
that although the results obtained with T7phia are up to the present in the highest 
degree encouraging, it is not possible to predict what the future may hold in store for 
it. No natural enemies of Tvphia of any importance are known, but in a foreign 
environment it is conceivable that hitherto unknown inimical influences may develop ; 
in consequence it would for the present appear unwise to relax the control at present 
practised and in point of fact more stringent control measures are in contemplation. 


The Cost of the Campaign. 

The total expenditure on the campaign against Phytalus smithi up to 31st December 
1919 amounted to Rs.232,100 from funds administered by the Department of 
Agriculture. Up to the year 1915, the money expended was provided entirely by 
Government; since then a system of contributions by estate owners has been 
inaugurated. During the year 1918-19 Government contributions amounted to 
Rs.24,000 and the estates’ contributions to Rs.8,500. Apart from the direct 


THE CAMPAIGN AGAINST PHYTALUS SMITHI, 169 


expenditure, a considerable sum of money has also been spent by estates in 
operations for beetle destruction, both in labour in the digging out of larvae from 
infested fields and in the provision of extra premiums, supervision and materials 
for the work of beetle collection. 

Throughout, the closest co-operation has been maintained with the planters 
concerned, meetings are periodically held for the purpose of discussing plans and 
details of the campaign with interested proprietors and managers. In this way 
harmonious and efficient work has been secured, and many obvious administrative 
difficulties have been successfully avoided or overcome. 

Very recently the planting body of the Colony have decided unanimously that, in 
view of the fact that the pest may be held to constitute a serious menace to the 
greater part of the sugar industry of the Colony, the entire cost of control should be 
borne by the planters ; and they have recommended to Government the imposition 
of an export tax of two cents of a rupee per hundred kilos of sugar exported for the 
purpose of defraying the cost of the campaign. 

The sum of money likely to be provided as the result of this measure will suffice 
for a further extension of activities, and, given the favourable results so far achieved, 
there is reason for hoping that within a comparatively short period the incidence of 
the pest may become reduced to comparatively small proportions. 

It is impossible to gauge exactly what would have been the effect if these measures 
of control had not been undertaken. It is, however, certain that the pest would 
have extended to a very much greater extent than at present, and in badly infested 
fields would have probably rendered cane-growing impracticable. The lower 
sugar-growing lands of the island, which provide a suitable environment for Phytalus, 
probably constitute two-thirds of the total cane-growing area; in the absence of 
rigid control measures it is probable that by now the beetle would have become 
endemic in the greater part of them. While therefore the exact loss which has been 
avoided cannot be estimated, one is safe in saying that it would certainly amount 
to several millions of rupees. On this account the measures taken and the 
expenditure incurred must be regarded as abundantly justified by the results. 


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Fig. 1. 


Fig. 2. 


Votes al. PART. 2; 


A night’s capture of Phytalus smithi. 


Destroying captured beetles by burning. 


PLATE 


a 


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= 
7 Zz =o 


Coe oe 
7 q 


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171 


NOTES ON INSECTS ACCIDENTALLY INTRODUCED INTO THE ISLAND 
OF MAURITIUS. 


By D d’Emmerez de CHarmoy, 


Government Entomologist. 


In respect of insect and, incidentally, fungous diseases of plants, Mauritius is 
remarkable in two respects ; the first being the very small number that are indigenous 
to the island, and the second the very large numbers that have been introduced. The 
desire to introduce exotic plants of a useful or ornamental nature is very widely 
spread in this Colony, and very extended plant introductions have taken place. 
Until comparatively recently no restrictions existed on such importations, and as 
a result insect and fungous pests in large numbers have become established to the 
sreat and permanent detriment of the major and minor agricultural industries of 
the Colony. For example, the cane-sugar industry was threatened with ruin in 
1842 as the result of the introduction of the borer, Proceras saccchariphaga, Bojer, 
from Ceylon ; at a somewhat later date great damage was done by the insect known 
locally as the “ Pou a Roche Blanche ” (Pulvinaria gasteralpha, Icery) ; and again 
quite recently the introduction of Phytalus smithi has proved a serious menace to 
the industry and has resulted in the loss of large sums of money. 

Minor industries have suffered in a corresponding degree. For example, the 
cultivation of coffee and vanilla has been in large part abandoned, partly as the 
result of the incidence of introduced cryptogamic diseases ; while in general it has 
been shown in another paper that out of 80 species of insects recorded as pests of 
minor crops in the Colony, 50 (or 62°5 per cent.) are of exotic origin. 

In relation to official measures of control, the first action taken was in the year 
1882, when an ordinance was passed dealing with the introduction of pests of the 
grape vine, the legislation being the outcome of a convention entered into by the 
local authorities with the Government of the Cape of Good Hope. 

Until 1910, no other protection was afforded ; in this year a second ordinance was 
passed empowering the Collector of Customs to destroy, or cause to be treated, plants 
and other articles, the importation of which was liable to lead to the introduction 
of pests and plant diseases. The ordinance remained inoperative until 1911, when, 
by proclamation, the powers thereunder were transferred to the Director of Forests. 

In 1913, the regulations under these ordinances were again revised, under Pro- 
clamation 81 of 1913, whereby the restrictions were made more comprehensive, 

and executive powers transferred to the Director of the then newly organised Depart- 
ment of Agriculture. 

Since that date a regular and efficient control has been exerted over plant importa- 
tions. The text of the Ordinance and the regulations made thereunder are given as 
an appendix to this paper. 

It is impossible, at the present time, to trace the manner in which many of the 
older insect pests of crops were introduced into the island, or even to give the exact 
date of their introduction. 


Wey D. D EMMEREZ DE CHARMOY. 


On the other hand, information regarding occurrences during the past twenty 
years is much more precise and a review of it is of interest, inasmuch as it affords 
indications of the channels of introduction and the harm that has been caused thereby. 


Phytalus smithi. . 

Though this beetle was detected in Pamplemousses only in the year 1912, there 
is no doubt that its introduction dates back to at least ten years before its existence 
was noted. The infested area, the large number of insects destroyed in the first year 
1912, the subsequent infestation of other localities, and its rate of spread in these 
localities, leave no doubt as to the time that the insect must have taken to spread 
over so large an extent, which in 1912 was about 1 mile in radius. 

Though it is impossible to make any definite statement, the probabilities are that 
the insect was originally introduced through the Royal Botanical Gardens, 
Pamplemousses, by means of cane plants imported from Barbados, packed in soil in 
Wardian cases. 


A Cecidomyiid Fly. 

This Cecidomyiid, which it has not been possible to identify up to now, and the 
origin of which cannot therefore be traced with certainty, was probably imported in 
the year 1909 with a lot of mango plants from India by a sugar estate proprietor of 
Grand Port. 

A few years later, the presence of this insect was revealed simultaneously at Port 
Louis and at Mahebourg, on mango trees, the leaves of which bore small conical 
galls grouped together irregularly and containing Cecidomyiid larvae. These 
localities were at first the only two affected, but the infestation was then so 
widespread that there was no hope of destroying the pest. After having remained 
localised in these two places until 1918, this fly was found at Grande Riviére, then at 
Riviére Noire, and can now be seen in almost all localities in the island, in varying 
numbers. 


Dacus d’emmerezi. | 

This Trypetid fly was described by Professor Bezzi in 1917 in the Bulletin of 
Entomological Research. It is not an indigenous species, as its presence was only 
detected in 1915, in the course of investigations concerning Dacus sygmoides. It 
would be difficult to say whence it came, though one might reasonably suppose that 
it has been introduced from Madagascar by means of cucurbitaceous fruits, as it 
would have been difficult in infested fruits coming from a more distant place to pre- 
serve the healthy appearance which alone could have warranted their safe transit. 

This species is nowadays far more common than Dacus sygymoides, and exhibits a 
partiality for marrows and pumpkins, constituting a pest in the cultivation of these 
vegetables. 


Echidnophaga gallinacea. 

This flea found its way into Mauritius on fowls imported from Vohemar in 1913, 
and others which were introduced from South Africa in 1914. The insect proved 
troublesome at Mapou during the early days of its introduction ; it has not spread 
to the other districts up to now. 


NOTES ON INSECTS INTRODUCED INTO MAURITIUS. 173 


Sarcopsylla penetrans. 

The jigger has been repeatedly introduced from Madagascar and Farquhar Island 
since 1910. Rare cases of infection were noticed at Port Louis several years ago. 
The flea has since never been found elsewhere. 


Solenopsis sp. (Red Ant). 

It has not been possible to identify this ant up to now, and it is not therefore 
possible to say what is its country of origin. It is stated to have existed in Pample- 
mousses for the last twenty years. The writer first encountered it at Rose Hill in 
the district of Plaines Wilhems some fifteen years ago, when it was only to be found 
in a few places. 

It is now widespread throughout the island, being quite common on the coast 
and in hot places, where it has become a regular pest ; it is still relatively rare in the 
higher districts, where it has not hitherto attracted special notice. It rarely 
establishes itself in houses ; on the other hand it is a serious pest of food crops 
wherever it has established itself, building its nests deeply in the soil and extending 
its subterranean galleries in every direction for long distances. It steals small 
seeds from seed beds and eats on the spot larger seeds which it cannot carry away ; 
it also attacks young seedlings. 


Coccidae. 

As one would expect, importations have been more numerous in this family 
than in any other, not only from the year 1900 but also previous to that date. 
In a monograph published by the author in 1900, 54 species were mentioned, among 
which three only could be considered as indigenous. This number has since been 
increased by eight more ; these are :—Asterolecanium spectabile, Newst., Aspidiotus 
destructor, Sign., Aspidrotus mauritianus, Newst., Chionaspis simplex, Green, 
Lecanium mangiferae, Green, Lecanwum hesperidum, L., Pulvinaria antigoni, Green, 
and Pulvinaria sp. 


Asterolecanium spectabile and Aspidiotus mauritianus. 

These scales were observed for the first time in 1915 in Pamplemousses Gardens 
and at the Botanic Gardens, Curepipe, on various species of palms and on Cycas 
revoluta and Cycas sp. They cause much harm to these plants in spite of the fact 
that the insects are highly parasitised by various Chalcids during a certain part of 
the year. 


Aspidiotus destructor. 

This species also appears to be of relatively recent introduction, as it is only 
quite lately that guavas, the plants on which they are most abundant, show visible 
signs of infestation. It is probable that the insect was imported from South Africa, 


whence a considerable number of plants have been introduced during the last ten 
years. 


Chionaspis simplex. 
This species has hitherto been found only on bamboos and is relatively rare, 


causing no apparent damage to this plant; it presents no importance whatever 
from an economic standpoint. 


174 D. D EMMEREZ DE CHARMOY. 


Lecanium mangiferae. 
The importation of this scale probably does not date back further than 1907, 


as it was only in 1908 that the first infested trees were recorded in Moka. It would 
be difficult to state its origin, as the author has observed it in Madagascar and in 
Réunion in 1917, where many trees, especially mangoes, were severely attacked. 
In this Colony, it is in Grand Port and especially in the town of Mahebourg that 
this insect causes most damage. Mangoes, bread-fruits, Connamomum zeylanicum 
(cannellier) and litchis are often infested to such a degree that they do not bear 
fruit for several years. In the north of the island, where it exists, its presence 
is not apparent, either on account of unfavourable climatic conditions, or owing 
to a small Chalcid which parasitises 1t intensely. 


Lecanium hesperidum. 
It seems curious that this species, the distribution of which is world-wide, should 


not have been found in Mauritius before 1900. The effects of the insect are so 
striking that it is inadmissible that 1t could have escaped the author’s notice at the 
time when this group of insects was being worked upon. It was introduced from 
the Seychelles or from Madagascar with oranges. On the higher plateaux its 
presence is hardly perceptible, but it 1s much in evidence along the coast, where 
limes, particularly, suffer severely from its attacks. 


Pulvinaria antigoni. 

This species was introduced some 10 years ago from the Seychelles ; it was first 
discovered in Pamplemousses Gardens and a little later at Phoenix on Solanum 
wildenit and on Aristolochia. It is far from beimg widespread in the island and is 
relatively common on these plants for only a few months of the year, being highly 
parasitised by two minute Chalcids. It is up to now of no economic importance. 


Gagrella feae. 

This Phalangid, which was imported from Burma some 10 years ago with cargos 
of rice from Rangoon, is a striking instance of the rapidity with which an exotic 
insect can spread itself, when conditions prove favourable to its development and 
when nothing intervenes to check its spread and multiplication. The first specimens 
were seen by the author in 1910 and had been captured in the town of Port Louis by 
Mr. Lauricourt Olivia, taxidermist to the Museum. Three years later it had reached 
Grand River, 7.e., 26 miles from the original point of occurrence, and in the course 
of the five following years, the other districts of the Island had all become more 
or less invaded ; nowadays it is to be found everywhere. It has established itself 
in all towns and has crept over to the forests, where it exists in considerable numbers. 
It is sometimes so common in houses as to constitute a regular domestic pest. 

These Phalangids shun light and have the habit of collecting in considerable 
numbers in dark places. They spread out rapidly whenever they are disturbed, 
but will come back soon afterwards. 


NOTES ON INSECTS INTRODUCED INTO MAURITIUS. 175 


Anoplodesmus saussurel. 

This millipede was introduced into the island about 1904-1905, in Wardian cases 
containing rubber plants imported from Ceylon by an estate situated in Vieux 
Grand Port district. One year later the writer was in a position to say with 
certainty that the creatures occupied already the whole mountain range that 
separates Grand Port district from Flacq, and that the western side was no less 
infested than the eastern side. 


Conclusion. 

It follows from the above that within the last 20 years, 14 exotic species of insects 
are known to have been accidentally introduced into this Colony. Of these 14 
species, two have no significance from an economic standpoint; the 12 others 
have already caused considerable damage. 

These facts demonstrate emphatically that the restrictions established through 
the Department of Agriculture in 1913, though they may hamper to some extent 
freedom of importation, fulfil an urgent requirement in protecting the agricultural 
products of this Colony, which in the past have suffered considerable losses that 
could have been avoided altogether by the adoption of the measures now in force. 


APPENDIX. 


ORDINANCE 4 oF 1910. 


99 


1. In this Ordinance “ article’? means any seeds, plants, cuttings, or any 
package, covering or thing that may have come directly or indirectly from any 
Country or place named in any Proclamation issued under this Ordinance. 

2.—(A). It shall be lawful for the Governor in Executive Council by Proclamation 
to prohibit the importation of any articles from any country or place named in such 
Proclamation which, in his opinion, are likely to be a means of introducing any 
plant disease from such country or place. 

(B). In like manner the Governor in Executive Council may prescribe the con- 
ditions under which alone the importation of any articles shall be permitted that 
may have come directly or indirectly from any country named in such Proclamation. 

3. Any article coming from any country or place the importation from which is 
prohibited, and any article arriving from a country or place the importation from 
which is permitted upon certain conditions only, until and unless such conditions 
shall have been complied with to the satisfaction of the Collector of Customs, shall 
be deemed to be prohibited goods within the meaning of the Customs Ordinance 
No. 28 of 1892; and any such conditions as aforesaid shall be deemed to be 
restrictions within the meaning of the said Ordinance. 

4. So long as any Proclamation as aforesaid is in force any articles mentioned 
therein coming from parts beyond the sea may be deemed to have come from a place 
the importation from which is prohibited, and may be treated accordingly, unless 
the importer satisfies the Collector of Customs to the contrary. 

(687) . F 


176 D. D EMMEREZ DE CHARMOY. 


Proclamation 81 of 1913 issued under the above Ordinance provides :— 

1. The importation into Mauritius from any country or place whatever, including 
the Dependencies of Mauritius, is absolutely prohibited of :— 

(a) grape-vine cuttings and plants, except when covered by a certificate from 
the Board of Agriculture (or other competent authority) of the country 
of origin that the vines have not been exposed to the infection of phylloxera. 
for the six weeks prior to the date of shipment ; 

(6) earth, and leaf and garden mould ; 

(c) sugar-canes or cuttings thereof, live plants of all sorts, including roots, 
tubers, cuttings and grafts in any description of earth or soil ; 

(d) dung or animal droppings (except guano) ; 

(e) forage ; 

(f) timber with the bark on. 

2.—(A). The following articles may not be introduced unless written permission 
has previously been obtained from the Director of the Department of Agriculture :— 

(a) sugar-canes or cuttings thereof ; 

(b) living plants or bulbils of Agave or Fourcroya ; 

(c) tea plants. 

(B). The issue of a permit shall be in the discretion of the Director of Agriculture, 
who may attach conditions to the permit and may limit the number of plants, cuttings 
and others aforesaid to be introduced. 

3. The following will be inspected at the port of entry :— 

(a) sugar-canes or cuttings thereof ; 

(b) live plants of all sorts, including roots, tubers, cuttings, grafts, and buds ; 

(c) fresh citrus fruits from all countries except the Dependencies of Mauritius. 

The consignee or his agent shall, upon request, open the coverings and afford the 
inspecting officer every facility for conducting the examination. 

4. If, on such inspection, the articles are found to be not free from pest or diseases, 
they may be ordered to be destroyed by the consignee or his agent under the 
supervision of the inspecting officer, or to be subjected to such process of disinfection 
or treatment as the inspecting officer may prescribe, and the consignee or his agent. 
shall pay in respect of such treatment such fees and charges as are prescribed. 

5. If the removal is authorised, all sugar-canes or cuttings thereof and live plants. 
of all sorts, including roots, cuttings, grafts, and buds, shall be planted in a nursery 
apart from growing plants of the same kind and shall be subject to inspection by the 
Department of Agriculture from time to time during twelve months from the date. 
of importation. 

_ 6. If, on such inspection, the articles are found to be not free from pests or diseases 

not known to occur in the Colony they shall be rooted out and destroyed by the owner 
under the immediate supervision of the inspecting officer, and if found to be not free 
from pests or diseases known to occur in the Colony, they shall be so treated by the 
owner as may be directed by the Director of the Department of Agriculture. 

7. Non-compliance with the conditions imposed in articles 4, 5 and 6 with respect. 
to the disinfection, treatment and planting of the articles imported will render them 
liable to destruction at the expense of the owner. 


NOTES ON INSECTS INTRODUCED INTO MAURITIUS. Rit 


8. Provided that nothing in the above shall prohibit or prevent the Government 
from making such importations and introductions of such live plants, cuttings and 
others aforesaid, as it may require for scientific investigations. 


The provision for prohibition of certain imports is to protect our staple industries 
from pests and diseases which could be introduced in shipments, and which would 
at the same time be very difficult to detect at the port of entry. 

The sugar-cane, fibre and tea industries are also protected against the introduction 
of pests and diseases by imports only being allowed from countries and localities 
where serious diseases are non-existent. The Department of Agriculture through 
its exchanges of scientific publications is well informed of the occurrence of pests 
and diseases in other countries, and therefore can afford assistance to the principal 
industries of the Colony by having control of the imports of certain living plants. 

Fumigation and disinfection of certain imports after inspection is provided for, 
in order to safeguard against the introduction of diseases and pests of a cosmopolitan 
character, and the provision for plantings to be made in nurseries is made so that 
insects or diseases which may by chance have passed through the port of entry 
inspections and treatment may be detected before they have become spread in the 
country. 

By Proclamation 52 of 1913 it is provided that the importation into Mauritius 
from any country or place whatever, including the Dependencies of Mauritius, 
of all living insects except such as may be required by the Government for scientific 
purposes is absolutely prohibited. 


(687) F2 


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179 


A PANNIER HOPPER-DOZER. 
By C. B. Wiuu414ams, MA, F.ES., 
Department of Agriculture, Trinidad. 
(Pratm V1). 


In the course of my work on the froghopper (Tomaspis saccharina, Dist.) damaging 
sugar-cane in Trinidad, it was found that the adult insects could be caught in very 
large numbers at the time of their broods by drawing large nets over the cane plants 
in the late evening and early morning. A number of different types of net were 
designed and experimented with, and the one described below has proved the most 
satisfactory up to the present. 

Unfortunately, owing to the very limited period of daylight during which the 
adults can be so caught, the use of these nets as a practical control has not so far 
been a success. The account below is given entirely in the hope that it may contain 
some new suggestions that might be applied or adapted to the control of other pests 
under other conditions. 

It was necessary to get a net of good width that could be moved rapidly over canes 
of various heights, without damaging them, and with the least expenditure of’ 
labour. These results were obtained by having two nets in the form of panniers. 
on the sides of an animal (in this case a mule), the whole being wide enough to sweep :: 
two rows of cane planted five feet apart when the animal walks between the rows. 

Very little description is necessary if the figure and plate are examined carefully. 
Most of the details are unimportant and could be varied to suit local conditions. 

Kach net is hinged behind and is supported in front by a stout cord or wire that 
is attached to the front supporting bar. This is loose in its bearings, and by rotating 
it the front of the net can be raised or lowered, which can be done by the boy in the 
saddle, without getting down, as the height of the canes changes. The nets in the 
machine figured could be lowered so that the bottom edge was only a few inches 
from the ground, when they could be used in long grass, or raised until there was 
about three feet clearance. It was not found feasible to send mules through canes 
requiring the nets higher than this. By removing the stop on the rear bar and 
unhooking the front supporting cord, each pannier can be removed in less than a 
minute. 

The nets are open in front and may be smeared inside with some sticky substance, 
or the opening could be fitted with a shutter arrangement of netting to prevent the 
escape of the insects. In the present machine they are of wire mosquito netting, 
but strong cloth or even thin wood might be used. The advantage of the net is 
that it allows the air to pass through, and does not create a back-draught. 

The framework of the nets and saddle was constructed of wooden bars, one inch 
by two inches, while the main supporting bars, rear and front, were one inch iron 
tubing. The saddle should be well padded and securely strapped down with two 
girths, 


180 C. B. WILLIAMS. 


Provided that the nets were not too close to the ground no difficulty was found 
in taking them over rough ground, open field drains or even small gullies. It need 
scarcely be added that it is as well to start work gently and with an unusually quiet 


animal. 


" REAR BAR. 


PROJECTS 


2’ 3" 


END 


Sd -- 


v 
2 


OJECTS 


Fig. 1. Details of pannier hopper-dozer. 


181 


A LIPARID MOTH (OCNEROGYIA AMANDA, STAUD.) DESTRUCTIVE 
TO FIGS IN MESOPOTAMIA. 
By P. A. Buxton, M.A., F.ES., 
Fellow of Trinity College, Cambridge. 


There are few countries in which agriculture is more important and the enemies 
of the crops less known than Mesopotamia. The following notes on a serious pest 
of the fig-tree in various parts of the Baghdad vilayet were put together on the 
River Diyala, north-east of Baghdad, at the end of July 1918. The Liparid moth 
which is so injurious has been determined by Sir G. F. Hampson as Ocnerogyia 
amanda, Staud. The larvae devoured all the fig leaves completely, with the 
exception of one or two of the large vascular bundles. The fruit shrivelled and 
dropped before it was ripe. 


Life-History. 

The eggs are easily found in patches of from twenty to a hundred, often laid 
so that they did not touch each other ; the groups of eggs are found on the lower 
parts of the trunks, more rarely on the underside of the lower leaves, and on rubbish 
on the ground. The female does not deposit her anal hairs upon the eggs. The 
eggs are large* for the size of the moth (horizontal diameter 1°39 mm., height 
1°23 mm.). 

As Dr. G. A. K. Marshall has pointed out to me, eggs large in proportion to the 
size of an insect are generally laid by species which either aestivate or hybernate 
in the egg stage. This is clearly not the case in the species that we are now con- 
sidering, which is continuous-brooded. 

The larvae are almost entirely night feeders, and conceal themselves before 
7.0 a.m. in summer among dead leaves beneath the fig-trees, in cracks in soil or 
in mud walls and similar situations. They may occasionally be found by day on, 
and especially under, the leaves in shady parts of the gardens. The hairs of this 
species produced no urticating effect on my skin, which is however by no means 
sensitive. 

The pupa is placed in cracks in soil, and under overhanging portions of mud walls. 

So far as I could detect during my short stay in the area in which I found this 
species, the two sexes fly with equal readiness if disturbed during the day-time ; 
they do not fly in sunshine unless disturbed. 

We have no knowledge of the time of year at which the pest begins to be trouble- 
some. It has been actually observed from the end of July to the end of September. 
It is apparently a species that breeds continuously during these months, as all the 
stages can be found at the same time. This is quite unusual, for the very great 
majority of Mesopotamian Lepidoptera aestivate throughout the summer, which 
is excessively hot ; we do not know whether they go through the summer as eggs 
or pupae. 


* Compare Orgyia antiqua, 0°82 by 0°68 mm. ; Liparis monacha, 1:045 by 0°77 mm. 


182 P. A. BUXTON. 


Distribution. 

In the year 1918 O. amanda was a very serious pest of figs at Ba‘quba (Bakubah) 
on the River Diyala, about thirty miles north-east of Baghdad ; also at Mendali, 
Shahroban, and Balad Ruz, all of which are within sixty miles of Ba‘quba towards 
the Persian frontier ; also at Kerbela on the west side of the River Euphrates, 
50 miles 8.8.W. of Baghdad. This last was important, because the large black 
figs of Kerbela are famous all over lower Mesopotamia and the whole crop was 
destroyed by O. amanda in 1918. So far as I know, figs, though widely grown, are 
not a principal crop in any part of the country. The incidence of the pest appears 
to be very sporadic ; I failed to find it at Khaniqin, about 60 miles up the River 
Diyala from Ba‘quba, or at Baghdad, Basrah, Qurnah, Nasiriyah, or Suq-ash- 
Shuyukh. In the year 1919, as I am informed by the Director of Agriculture, 
Baghdad, there were no reports of its incidence in Mesopotamia, and in the same 
year I failed to find it in North-West Persia at Qazvin and Enzeli, and at Tiflis in 
Transcaucasia. 

The larva is known to the Arab cultivators as Shimbaran, but the word is not 
in very general use. 

Preventive Measures. 

The habits of the insect lead us to believe that it should be easy to keep in check, 
and the following measures may be mentioned :—(1). Arab cultivators burn the 
dead leaves and other rubbish in heaps beneath the trees. This destroys all stages 
of the insect, and also its hiding places. The damage done to the tree is shght. 
(2). Hand-picking the larvae from beneath the leaves shortly after dawn should 
be effective ; batches of eggs would be found at the same time. (3). Grease banding 
would no doubt catch the larvae on their morning and evening migrations, but 
it is unlikely that so expensive a method will be used when burning the rubbish 
is so efficacious. 


Description of the Moth. 

When Staudinger described the monotypical genus Ocnerogyia he had only the 
male of the present species before him ; the female has not previously been captured. 
Sir George Hampson has been good enough to redescribe the species as follows :— 

Family LrpaRipae. 
Genus Ocnerogyia. 

“ Ocnerogyva, Staud., Iris, iv, p. 254 (1891). Proboscis absent ; palpi very short, 
porrect, clothed with rough hair ; frons smooth, with large tuft of hair ; eyes rather 
small and elliptical, smooth; antennae of male bipectinate with long branches 
to apex, of female with rather shorter branches ; thorax and abdomen clothed with 
rough hair and without crests ; tibiae and tarsi moderately fringed with hair, the 
mid and hind tibiae with terminal pair of spurs. Forewing broad, the costa evenly 
arched, the apex rounded, the termen obliquely curved and not crenulate ; vein 
2 from beyond middle of cell ; 3 from well before angle of cell; 5 from above angle ; 
6 from upper angle; 7, 8, 9 stalked; 10, 11 from cell. Hind wing with the cell 
long ; vein 2 from beyond middle of cell ; 3 from well before angle of cell; 5 from 
just above angle; 6, 7 from upper angle; 8 connected with the cell by a slight 
oblique bar just before middle. 


A LIPARID MOTH DESTRUCTIVE TO FIGS IN MESOPOTAMIA, 183 


Ocnerogyia amanda. 

Ocnerogyva amanda, Staud., Iris, iv, p. 254 (1891). 

“ g. Head and thorax grey, tinged with reddish brown the lower part of frons 

and palpi yellowish ; abdomen fulvous orange. Forewing grey, tinged with reddish 
brown and urorated with rather elongate pale griseous scales; a rather diffused 
blackish discoidal bar. Hind wing uniform fulvous orange, the cilia brownish grey. 
Underside of forewing without dark irroration or discoidal bar, a small rather 
diffused fulvous orange patch below middle of cell ; hind wing with the costal edge 
brownish grey. 
_ “ O. Antennae with the shaft whitish ; abdomen pale grey tinged with brown, 
the basal segments and sides tinged with yellowish ; hind wing pale grey, tinged 
with brown, the inner area with some pale fulvous hair; underside of forewing 
without the fulvous orange patch below the cell. 

Ezp., 3 38, 29 40 mm. 

Upper Syria: Mardin. Mesopotamia: R. Diyala, Ba‘quba (P. A. Buxton), 1 3, 
1 Qin Brit. Mus.” 

Description of the Larva. 

The larva presents points of difference from that of other Liparrpaz, and I have 
drawn up the following description from spirit specimens in the last instar. 
Unfortunately I was engaged with other work at the time when I found the larvae, 
at the height of the Mesopotamian summer ; my notes only say that the skin was 
grey-green in colour, and that the intersegmental membrane was neither paler 
nor darker than the rest. The setae were silky and almost colourless. The general 
appearance is shown in text-fig. 1 (A & B) and suggests an Arctiid rather than a 
Liparid. There are none of the dense brushes of hair with which we are so familiar 
in Orgyia and Dasychira. The larva lacks also what Fracker calls “‘ clavate-plumed 
setae.” The colourless setae are very finely and evenly serrate. It will be noticed 
that long setae arise from the prothorax (actually from a large verruca formed by 
the fusion of Kappa and Rho), and curve forwards over the head of the larva ; 
these setae are more than half the length of the larva, but in fig. 1.B only their 
basal half is shown. On the meso- and metathorax and abdomen the longest 
setae grow out sideways on verrucae Kappa and Pai. Among the silky setae a 
few brown ones (b.s.) may with difficulty be found. They are always long and 
thick, and arise from the large verruca (g.t.) (Rho plus Kappa) of the prothorax 
in some numbers, and as single setae on Beta of the meso-thorax, and Rho on 
abdominal segments 1-8 inclusive. 

Chaetotaxy. The head. All the primary setae of Forbes can be easily seen 
except 11, v, and vi. The last two are probably present but cannot be distinguished 
with certainty among the rather large secondary setae which are sparsely scattered 
over the lower and more lateral parts of the head. There is only one secondary 
seta on the adfrontal, situated below adfrontal i; this is fewer than is usual among 
the Lipartpak (Forbes). 

The thorax.* On the anterior margin of the prothorax, near the mid-dorsal line 
is a small group of setae, alpha, beneath it a still smaller group of setae, two or three 


ae describing the chaetotaxy of the thorax and abdomen [ have followed Fracker’s 
system. 


184 P. A. BUXTON. 


only, gamma. Beta and delta appear to be absent. In front of the large pro- 
thoracic spiracle an immense tubercle (g.t.) arises, pomting forwards. It is clothed 
with setae of both types, the brown and the colourless, which are more than half 
as long as the entire larva, and are directed forwards over the head. This verruca 
is very well developed in various Liparid larvae which I have examined, and is 
formed by the fusion of Rho and Kappa. This we see when we compare the pro- 
thorax with either of the succeeding segments, in both of which Rho and Kappa are 
separate. The process of fusion of these two verrucae, so complete on the pro- 
thorax, can actually be watched taking place on successive segments of the abdomen. 
Beneath this very large verruca is a second one, which consists of Pai with its asso- 
ciated setae ; this is recognisable in all segments to the ninth abdominal inclusive. 
Its setae project outwards and downwards and form a conspicuous tuft over the leg 
on all leg-bearing segments. 

Meso- and metathorax. The chaetotaxy is sufficiently shown in text-fig. 1 (B). 
Rho and Kappa are not fused. 


Abdomen. Beta (a, plus 3, plus secondary setae) can be seen on each of the 
first nine segments and is largest on the eight and ninth. The gradual migration 
of Kappa to a position behind but distinctly above the spiracle is interesting. It 
never coalesces completely with Rho except on the ninth segment, but the two 
groups, Rho and Kappa, are borne on one large partly divided verruca on all the 
abdominal segments. The group which represents sigma consists of many setae, 
and is borne on a definite verruca on the legless segments (1, 2, 7, 8, and 9) ; on the 
other segments it is only to be found with difficulty. No trace can be found of nu, 
tau, or mu ; we presume therefore that they are components of Pai. 

The legs and prolegs are covered sparsely with secondary setae which are believed 
to be of no phylogenetic significance. The prolegs (A and C, pl.) are long and armed 
with long crochets all of one size arranged in a mesoseries, that is to say a curved 
line concave outwards. On the outer side of all prolegs except the anal pair is a 
chitinous plate bearing secondary setae. The dorsal glands on abdominal segments 
6 and 7 are small ; the diameter of the mouth of the gland is less than verruca Beta ; 
these glands are believed to be characteristic of all LiparIDAE. 


The points of interest in the structure of the larva are :-— 

(1) Liparid larvae fall into two main groups: Ocnerogyia amanda falls into the 
first, many of which are densely hairy ike Arcrmpar. ‘The second group contains 
bizarre species armed with dense brushes (verricules) and clavate plumed setae ; 
Orgyiais a good and familiar example of this group. 

(2) Fracker states that in these insects there are “‘ three verrucae above the Kappa 
group on the mesothorax and metathorax.”’ Forbes says (p. 103) “ five warts on 
mesothorax.” (This comes to the same thing as beneath Kappa lies Pai). Ocnerogyra 
amanda forms an exception to these general statements ; as will be seen from text- 
fig. C, there are only four verrucae on the mesothorax, and metathorax ; two of 
which are above Kappa. 

(3) The eversible gland on the mesodorsum of abdominal segments 6 and 7 
appears to be the most constant characteristic of the larvae of LiparipaE. So far 
as we know, it is always present, though it may be very small. 


A LIPARID MOTH DESTRUCTIVE TO FIGS IN MESOPOTAMIA. 185 


x 
i 
\) 


— 


\ 
\ 
Ny 
( 


bs. 
\ p 


B. 
Fig. 1. Structure of larva of Ocnerogyia amanda, Staud. 


A. Abdominal segments 5-10 : 

B. Head, iiavasohid abd. segt. i | Magnified by about 35. 

In these figures certain segments are shown armed with their setae; others are 
denuded of setae and show the chaetotaxy diagrammatically. The long setae curving 
over the head are drawn about half their full Jength. The head is shown in outline 
only, without setae, eyes or mouth-parts. On I alpha and gamma are indicated, but 
are not lettered in order to avoid overcrowding the figure. 

C. Proleg of 3rd abdominal segment viewed from side, magnified by 150. 

I, II &III. Pro-, meso-, and metathorax ; 1-10, abdominal segments ; a.c, anal clasper ; 
b.s, large brown seta ; cr, crochets arranged in a line concave outwards, all of one size except 
those at the ends ; g.t, great tubercle of I; pl, proleg ; pt, plate on lateral side of pro- 
leg, bearing secondary setae ; sp, spiracle ; s.s, secondary setae ; B, Beta bearing alpha 
plus beta; on I Beta is figured but not lettered to avoid confusion ; K, the compound 
verruca Kappa; 11, large compound verruca above the insertion of leg or proleg ; 
P, aan bearing epsilon plus rho, and fusing with Kappa on I to form g.t. and 
partly fusing with it on 1-9; ¢, the nearly midventral sigma, which in this species is 
represented ‘by a tuft of setae. 

In fig. 1. A & B the compound verruca I has been marked 7- in error. 


186 P. A. BUXTON. 


References. 
W. T. M. Fores (1910). A Structural Study of some Caterpillars —Ann. Ent. 
Soc. Amer. ili, pp. 94-132, 11 plates. 
S. B. Fracksr (1915). The Classification of Lepidopterous Larvae.—lIllinois 
Biolog. Monographs, ui, no. 1, pp. 1-170, 10 plates. 
O. StaupINGER (1891). Inis, iv, p. 254. 


PLATE VI. 


PART 2. 


PGF 


VOE, 


ENT. RESEARCH. 


BULL, 


A pannier hopperdozer for catching insects on sugar-cane. 


Os 


aw 
_ 
re 
7 
be ~ 


: 


oe Fs 
omy nts 7 6 ee 


ie nian 


mm ly ‘ . 
we et lal Fe el 
Ws : - 


Po elected ae fase 


i: oe a3 cea oe Mat 


: end a. 


187 


COLLECTIONS RECEIVED. 


The following collections were received by the Imperial Bureau of Entomology 
between Ist April and 30th June, 1920, and the thanks of the Managing Committee 
are tendered to the contributors for their kind assistance :— 

Mr. E. Ballard, Government Entomologist, Madras :—120 Tabanidae and 69 
Tachinidae ; from South India. 

Capt. P. J. Barraud, Entomologist to the Egyptian Expeditionary Force :—79 
Culicidae, 22 other Diptera, 18 Hymenoptera, 141 Coleoptera, 6 Planipennia, 18 
Lepidoptera, 23 Rhynchota, 6 Orthoptera and 92 Odonata ; from Palestine. 

Mr. ©. F. C. Beeson, Forest Zoologist :—143 Curculionidae ; from India . 

Mr. John R. Bovell, Superintendent of Agriculture :—5 Diptera, 16 Hymenoptera, 
5 Thysanoptera, 1 species of Coccidae, 6 Aphididae, 6 other Rhynchota, and 5 Acari ; 
from Barbados. 

Major J. E. Boyd :—4 Tachinidae ; from Sandwich, Kent. 

Mr. J. B. Corporaal, Entomologist, Algemeen Proefstation, Medan :—12 Diptera, 
38 Coleoptera, and 4 Rhynchota ; from Sumatra. 

Mr. J. Sydney Dash, Director, Station Agronomique de la Guadeloupe :— 9 
Diaprepes beetles and 100 Aphididae ; from Guadeloupe. 

Division of Entomology, Pretoria :—11 Rhynchota and 21 Orthoptera; from 
South Africa. 

Dr. D. Duff :—5 Phlebotomus ; from Winnebah, Gold Coast. 

Dr. Eustace W. Ferguson :—5 Tabanidae, 4 Hippoboscidae, and 41 other Diptera ; 
from Palestine : 81 Tabanidae ; from Australia. 

Dr. Goldberg, Jewish Health Bureau :—1 Culicoides, 6 Culicidae and 1 Lyperosia ; 
from Palestine. 

Dr. Lewis H. Gough, Government Entomologist :—14 specimens of the butterfly 
Catochrysops eleusis ; from Egpyt. 

Hawaiian Sugar Planter’s Association :—10 Curculionidae ; from Honolulu. 

Dr. J. F. Illingworth :—906 Coleoptera and 8 Rhynchota ; from Queensland. 

Imperial Institute :— 10 Hymenoptera ; from Colombia. 

Dr. A. Ingram :—11 Culicidae, 9 Dipterous larvae, 238 Fleas, 7 Hemimerus, 
19 Anoplura, 3 Mites, and 73 Ticks ; from the Gold Coast. 

Mr. H. H. King, Government Entomologist :—2 Tabanus, 7 other Diptera, 47 
Hymenoptera, 26 Coleoptera, 106 Rhynchota, and 10 Crickets ; from the Anglo- 
Egyptian Sudan. 

Dr. W. A. Lamborn:—3,000 Fleas, 60 Chalcids, 317 parasitic Hymenoptera, 
145 other Hymenoptera, 9 Thrips, 116 Cimicidae, a number of Aphids, 10 other 
Rhynchota, 35 Anoplura, 104 Mallophaga, 983 Mites, 1 Glossina, 3 Auchmeromyia, 
282 other Diptera, 270 Dipterous pupae, 155 Ants, 1 Coleopteron, and 1 Lepidop- 
teron; from East Africa. 


Dr. J. W. Scott Macfie :—20 Phlebotomus, a collection of early stages of Pericoma, 


a number of other Diptera and Ants, and 12 other Hymenoptera ; from the Gold 
Coast. 


188 COLLECTIONS RECEIVED. 


Prof. G. H. F. Nuttall, F.R.S. :—2 Hippoboscidae ; from the Belgian Congo. 

Dr. H. Schouteden :—9 Hymenoptera, 12 Coleoptera, 23 Moths, and 1 Tettigoniid ; 
from the Belgian Congo. 

Mr. F. V. Theobald :—7 Chalcids (Pteromalus egregius) bred from the Brown- 
tail Moth : from Romney Marsh, Kent. 

Dr. B. Uvarov :—A collection of Phlebotomus ; from the Caucasus. 

Mr. Robert Veitch :—3 Tabanus, 5 other Diptera, 179 Coleoptera, and 23 
Microlepidoptera ; from Fiji: 7 Diptera, 20 Hymenoptera, and 31 Coleptera ; from 
New Zealand. 

Mr. C. B. Williams :—14 specimens of the Scolytid beetle Xyleborus grenadensis ; 
from Trinidad. 

Mr. R. C. Wood :—8 Tabanidae, 91 other Diptera, 298 Coleoptera, and 34 Rhyn- 
chota ; from Nyasaland. 


- IMPERIAL BUREAU OF ENTOMOLOGY. 
‘HONORARY COMMITTEE OF MANAGEMENT. 


< 


THE VISCOUNT HAROOURT, Chairman, 

~ Lirur.-Cotoven A. ALCOCK, €.LE., E.R.S. | hs Ble 

Mason BE. E. AUSTEN, DS.0. | ae icger 

Dr. A. G. BAGSHAWE, C.M.G aaa eh 

Masor-GEneraL Siz JOHN R. BRADFORD. K, ©. M. G., F Re = 

Masor-GENERAL Siz DAVID BRUCE, sme tine ae ae oe he ee 

Mr. J. C. FE. FRYER. 

Sm 8, F, HARMER, KBE. ERS. 

Pror. H. MAXWELL LEFROY.: 

‘Hon. E. LUCAS, 

Dr. R. STEWART MACDOUGALL, 

Str JOHN McFADYEAN: — 

Siz PATRICK MANSON, G.C.M.G., F. R.S. 
’ Sr DANIEL MORRIS, K.C.M.G. 

Pror. R. NEWSTEAD, F.R.S. 

Pror. G. H. F. NUTTALL, F.RB.S. 
‘Prov. E. B. POULTON, FRS. = | WN eds eS 
Lrevt.-Cotonen Sir DAVID PRAIN, ona, CLE, BRS. ag, a 
Sin H. J. READ, K.OMG., CB. | 3 cape. 
_ Taz How. N.C. ROTHSCHILD. | i 
Dr. HUGH SCOTT. ; 
Sm A. E, SHIPLEY, G.B.E., FERS. 
~ Mr. R. A. C. SPERLING, O.M.G.. ; | 
Sm. STEWART STOCKMAN. . - ce Pee ee eee 
| Mz FV. THEOBALD, tek as as ee a 

| 


) apa 
by ie oes pre the to, 
be OS ye eT eae. 


Me. C. race ae 


Director. 
od GUY- A. EK. MARSHALL, OM. 


ts 


iy | ‘Assistant Director, 
es - Dr. 5 A. NEAVE, 
Secretary, e 
ate, . 0. C, "PARKINSON, 


Ar ey rae ‘ Lf: ¢ ty 


189 
SOME NEW WEST INDIAN SPECIES OF THE MELOLONTHID 
GENUS LACHNOSTERNA. 
By GILBERT J. ARROW. 


Twenty-four West Indian species of this genus were enumerated in Messrs. Leng 
and Mutchler’s list of 1914, in addition to which four species from Porto Rico have 
since been named by Mr. Eugene Smyth (Journ. Dept. Agric. Porto Rico, i, 1917). 
Two other species have been assigned to the genus Phytalus, which is distinguished 
from Lachnosterna solely by the claw-tooth making an acute instead of a wide angle 
with the tip. As H. W. Bates found in his attempt to distribute the Central 
American species, it is impossible to make this the basis of a natural division, 
especially as in certain species (as in L. dilemma here described) it is confined to the 
male. 

The genus Phytalus was constituted by Erichson in 1847 for American and 
Oriental species, but was restricted to the former by Blanchard in 1850. Blanchard 
formed for Oriental representatives a new genus Brahmina, but without comparing 
the characters of the two. Brenske in 1892 confirmed this geographical division 
and set forth characters by which he maintained that a purely American genus 
Phytalus could be separated from the Eastern genera Brahmina, Holotrichia and 
others. These characters he defined as—(a) cleft claws, the inner branch being 
as long as or longer than the outer; (0) elongate antennal club in the male; 
(c) hollowed abdomen in the male ; the opposite conditions denoting the Oriental 
genus Brahmina. 

But Brenske’s statement of the case really points to the opposite conclusion to 
that drawn by him, for the application of his tests relegates the West Indian species 
Phytalus smithi, Arrow, and the Mexican P. omiltemus, Bates, to the genus Brahmina, 
while the Mexican P. platyrrhanus, Bates, combines the characters of the two genera, 
the inner branch of the claw being much shorter than the outer. As just mentioned, 
however, the cleft claw may be only a feature of the male sex, so that no single 
character has yet been found which is sufficient for the recognition of the female 
Phytalus, and those supposed to characterise the male are quite at variance with 
the present conception of the genus. In my opinion Phytalus, Brahmina and 
Holotrichia are quite undefinable, and I can see no alternative to treating them as 
one with Lachnosterna. 

In recent American works on this group of beetles the generic name Phyllophaga 
has been adopted. This name was introduced by Harris in 1826, eleven years 
earlier than the date of the characterisation by Hope of Holotrichia and Lachnosterna. 
It is however a nomen nudum, accompanied by no description, and is therefore 
upon the same footing as the numerous Catalogue names printed by dealers and 
others, to which obviously no scientific value can be attached. 

Including the five here described, thirty-five species of this genus are now known 
from the West Indian Islands, and many of them are probably of economic 
importance. 

(713) Wt.P2/164. 1,000. 12.20. B.&F.,Ltd: Gp.11/14 A 


190 GILBERT J. ARROW. 


Lachnosterna jamaicensis, sp. nov. (fig. 1, c). 

Castaneo-rufa, nitida, corpore subtus cum pygidio pallide flavo; oblonga, 
cylindrica, capite fortiter dense punctato, clypeo bilobato ; pronoto sat profunde 
haud crebre, medio paulo parcius, punctato, lateribus medio angulatis, postice 
fere parallelis, antice convergentibus, vix arcuatis, haud crenatis, angulis anticis 
fere rectis, posticis obtusis ; scutello utrmque minute punctato, medio fere laevi ; 
elytris modice fortiter et crebre punctatis, costa suturali duabusque discoidalibus 
vix perspicuis ; pygidio sat fortiter punctato ;. pectore dense flavo-villoso ; antennis 
9-articulatis, clava triphylla. 

6, Antennarum articulo 6° breviter laminato, clava modice elongata ; tarsis 
posticis quam tibius paulo longioribus: 9, antennarum clava brevi, tarsis posticis 
quam tibiis brevioribus. 

Long. 17-18 mm. ; lat. max. 8°5-9°5 mm. 

Jamatoa (A. H. Retchie). 

It is a rather narrowly elongate insect, of a mahogany-red colour, with the lower 
surface and pygidium pale yellow. The entire upper surface is shining, but rather 
strongly punctured, the head densely so. The sides of the prothorax are not crenate 
and are almost straight from the middle to the front and hind angles. 

The description of L. cylindrica, Burm., the exact habitat of which is unknown, 
almost applies to this species, but, although the joint preceding the antennal club 
of the male has a long sharp process as there described, there is not an abbreviated 
lamella to the club in addition, nor is the pygidium almost smooth. 


Fig. 1. Dorsal and lateral aspects of aedeagus of male 

of (a) Lachnosterna dilemma, sp.n.; (b) L. trinitatis, 

Arrow ; (¢) L. jamaicensis, sp. n. ; (d) L. antiguae, sp.n. ; 

(e) L. montserratensis, sp.n.; (f) L. a@emosa, sp.n. 

Claws of (k) L. montserratensis; (1) L. dilemma, 3; 
(m) L. dilemma, 9. 

The only species of the genus hitherto recorded from Jamaica is L. Jervida, 
Fabricius, which, although said by its author to be North American, Is attributed 
by Burmeister, who examined the type, to Jamaica and San Domin go. That 
species has the elytra pubescent and pruinose, while in the new species they are 


shining and quite devoid of hair. 


SPECIES OF THE MELOLONTHID GENUS LACHNOSTERNA. 191 


Lachnosterna montserratensis, sp. nov. (fig. 1, e, 4). 

Castaneo-rufa, nitida, elytris abdomineque (cum pygidio) leviter pruinosis, illis 
parcissime setosis, metasterno dense haud longe flavo-villoso ; oblonga, cylindrica, 
capite lato, fortiter sat dense aequaliter punctato, clypeo valde bilobato, vix excavato; 
pronoto aequaliter fortiter punctato, lateribus leviter crenatis, angulis omnibus 
obtusis, posticis rotundatis ; scutello fortiter punctato, medio fere laevi; elytris 
modice fortiter et crebre punctatis, costa suturali duabusque discoidalibus vix 
perspicuis ; pygidio fortiter punctato ; pedibus gracilibus, unguibus valde dentatis, 
dente reclinato ; antennis 9-articulatis, clava triphylla. 

3, Clava antennali modice elongata, pedum posticorum calcaribus longis, tarsis 
quam tibiis longioribus. 

Long. 17°5-19 mm. ; lat. max. 9-10 mm. 

MONTSERRAT. 

In size, colour, sculpture and general appearance this rather closely resembles 
L. jamarcensis, but there are numerous points of difference. There is a rather 
faint bloom upon the elytra, and this is more apparent upon the pygidium and 
abdomen. These are also more reddish in colour than in the Jamaican species. 
The head and pronotum are shining and strongly punctured, as in the latter, but 
the head is broader, the eyes are larger, the clypeus flatter and more strongly 
-bilobed. The punctures of the pronotum are rather more numerous, the lateral 
margins are slightly crenate and the hind angles rounded. 

I have not seen a female of this species. In the male the 6th joint of the antenna 
is not produced as in that of L. jamaicensis. The claw-tooth is very strong and 
sharp, inclined a little backwards and separated by a yery acute notch from the basal 
dilatation. 


Lachnosterna antiguae, sp. nov. (fig. 1, d). 

Testaceo-rufa, corpore subtus, pedibus antennisque flavis; cylindrica, parum 
elongata, pruinosa, haud nitida, pectore dense abdominisque medio parce flavo- 
hirto ; capite haud lato, grosse et rugose punctato, clypeo nitido, excavato, bilobato ; 
pronoto sat fortiter haud crebre punctato, lato, lateribus fortiter arcuatis, vix 
crenulatis, angulis anticis obtusis, posticis rotundatis ; scutello inaequaliter punc- 
tato; elytris leviter modice crebre punctatis, costa suturali duabusque discoid- 
alibus vix perspicuis ; pygidio paulo densius punctato ; pedibus modice gracilibus, 
tibiis anticis tridentatis, unguibus medio fortiter dentato ; antennis 9-articulatis, 
clava triphylla. 

d, Antennarum clava ad stipitem longitudine aequali, articulo 6° brevissime 
laminato, pedum posticorum tarsis quam tibiis longioribus; Q, clava antennali 
brevi, pedum posticorum tarsis quam tibiis multo brevioribus. 

Long. 14-17 mm. ; lat. max. 7-9 mm. 

Anticua (H. A. Tempany—April; H. A. Ballou—March; E. Elridge—July). 

Dominica (H. A. Ballou—May). 

Beetles and pupae were found in numbers in the soil of the cane-fields in March. 

The beetle is testaceous red in colour, its upper surface covered with a dull bloom 
and the clypeus alone shining (in fresh specimens). The clypeus is rather narrow 

(713) 7 a2 


192 GILBERT J. ARROW. 


and distinctly bilobed, the forehead is rugosely punctured, the pronotum strongly 
and irregularly, the elytra more finely and closely punctured, and the pygidium still 
more so. 

There is considerable resemblance to L. denticulata, Bl., from Martinique, of which 
M. Lesne has kindly sent me a co-type for comparison, but the new species is less 
narrowly elongate, the head is relatively narrower, the forehead more coarsely and 
rugosely punctured, and the sides of the prothorax are less distinctly crenulated. 
In size, colour and general appearance it also resembles the Mexican L. anodentata, 
Bates, but it is not hairy above. It is also readily distinguishable by the bilobed 
clypeus. 

In the male the club of the antenna is as long as the entire footstalk, in the female 
about as long as the scape. In the latter the hind tarsi are very short. 


Lachnosterna acinosa, sp. nov. (fig. 1, /). 

Brunnea vel rufo-brunnea, capite et pronoto fuscis, breviter griseo-setosa, pruinosa, 
sed capite, pronoti parte antica, pygidio pedibusque nitidis, metasterno dense flavo- 
villoso ; ovalis, convexa, capite haud lato, clypeo brevi, crebre punctato, excavato, 
margine vix exciso, fronte grosse et rugose punctata, tumida; pronoto grosse 
inaequaliter punctato, punctis setis erectis brevibus et longis intermixtis instructis, 
lateribus medio angulatis, vix crenulatis, haud arcuatis, angulis anticis et posticis 
fere rectis, anticis autem leviter acuminatis; scutello elytrisque minute et crebre 
punctatis, costa suturali aliaque discoidali vix perspicua ; pygidio nitido, subrugose 
punctato; antennis 10-articulatis, clava flava; pedibus utriusque sexus sat 
gracilibus. 

dg, Abdomine medio excavato, segmento penultimo ventrali medio dense spinoso. 

Long. 17-19 mm. ; lat. max. 9-10 mm. 

TRINIDAD (Ff. W. Urich). 

This is closely related to the Mexican species L. longipilosa and misteca, of Bates, 
differing from them most obviously in having the pygidium shining, instead of prui- 
nose. As in those two species the upper surface has both a bluish bloom and a_ 
clothing of fine setae, but the bloom is absent from the head and the anterior part 
of the pronotum. In the male sex the group is also characterised by the hollowed 
ventral surface of the abdomen and a peculiar spine-studded area in the middle 
of the penultimate segment. 

The head is rather small, the clypeus scarcely notched, and the forehead rugosely 
punctured. The pronotum is irregularly punctured and bears moderately long 
erect hairs intermixed with shorter setae. The elytra are rather finely and closely, 
the pygidium more coarsely, punctured. The antennae are 10-jointed and, like 
the slender legs, differ little in the two sexes. 


Lachnosterna dilemma, sp. nov. (fig. 1, a, 1, m). 

Nigro-picea, corpore subtus paulo rufescentiore, ubique subtiliter dense griseo- 
sericea ; ovata, convexa, creberrime punctulata, capite haud magno, grosse et 
confluenter punctato, clypeo brevi, margine rotundato, reflexo; pronoto dense 
minute punctato, angulis anticis paulo acutis, posticis obtusis; scutello paulo 
minus dense punctato ; elytris aequaliter dense punctatis, postice setis erectis parce 


SPECIES OF THE MELOLONTHID GENUS LACHNOSTERNA. 193 


instructis, costa suturali angusta duabusque discoidalibus parum perspicuis ; 
abdomine linea piligera circumdato, pygidio dense punctato et breviter erecte flavo- 
pubescente, metasterno dense haud longe pallido-villoso, pedibus modice gracilibus, 
antennis 10-articulatis, clava triphylla, flava. 

3g, Unguibus anguste fissis; 9, unguibus fortiter dentatis (vel apicibus late 
divergentibus). 

Long. 20-22 mm. ; lat. max. 10°5-12 mm. 

TRINIDAD: Verdant Vale (f. W. Urich—June). 

In size and general appearance this is rather like L. trinctatis, Arrow (fig. 1, 6) from 
the same island, but L. dilemma is more oval in shape, much more finely and closely 
punctured, especially upon the pronotum, and clothed everywhere with a very fine 
and close silky grey pubescence. Around the abdomen, at the line of contact with 
the outer edges of the elytra, the pubescence abruptly ceases and a fringe of long 
hairs is found. This, so far as I know, does not occur in any allied species. 

The abdomen of the male is hollowed beneath and the last ventral segment is 
concave, smooth and shining, whilst that of the female is convex and strongly 
punctured. ‘The hind tarsi of the female are little shorter than those of the male, 
but the tibiae are more dilated at the end and the spurs much shorter and broader. 
The claws of the male are narrowly cleft, but in the female the two branches diverge 
widely. 3 

There is considerable resemblance to the Central American ZL. setifera, Burm., 
but the clothing of the upper surface is finer, the clypeus a little shorter and the 
angles of the prothorax are less sharp. The male is easily distinguishable by its 
narrowly cleft claws and the normal straight suture between the last two ventral 
segments. There is also a resemblance to L. (Phytalus) cometes, Bates, but that is 
lighter red in colour, rather more elongate, and the abdomen has no line of 
demarcation between the dorsal and ventral parts corresponding with the outer 
edges of the elytra. | 


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195 


NOTES ON BLOOD-SUCKING FLIES IN NORTH RUSSIA DURING THE 
SUMMER OF 1919. 


By Major A. D. Fraser, D.8.0., M.C., R.A.M.C. 


With an Appendix containing a List of Species, by Major EF. E. Austen, D.S.O. 


Locality. 

Most of the following observations were made in the vicinity of Obozerskaya, 
a small village on the Archangel-Vologda Railway, 474 versts from Vologda or 
roughly 80 miles due south of Archangel. I arrived at Obozerskaya and reported 
to the Headquarters of the Vologda Force on 14th June 1919, a few days after 
disembarking at Archangel. 

Further experience and a few of the specimens collected were obtained during 
tours of inspection on the Onega River Front in the last week of June, and on the 
Seletskoe Front early in August. 


Nature of Country. 

The country in the whole of this district is one continuous forest of pies, firs, 
etc., and although there is comparatively little undergrowth, any movement through 
the forest other than along the recognised tracks is rendered extremely difficult 
on account of the swampy nature of the ground, which remained waterlogged 
even in July, when the heat was quite trying. 

Excepting those along the railway, most of the villages are situated on the banks 
of rivers or lakes, and around the majority small clearings are cultivated or used 
for grazing. At Obozerskaya the clearing was very small, and there was no culti- 
vation ; the nearest open water was a lake about two miles from the village. Along 
one or two of the larger rivers grazing land is more plentiful, and fair-sized herds of 
cattle are to be seen. 

Remarkably few birds or animals of any kind are met with in the forest, though 
ants are numerous everywhere. 


Tabanidae. 

Before I arrived in the country mosquitos were numerous and troublesome, and 
the troops had been provided with mosquito-nets and veils. Midges were also 
annoying, but otherwise no biting flies were in evidence until 20th June, a warm 
sunny day following a couple of days of rain, when numbers of TABANIDAE appeared. 
A day or two later they had increased to such an extent that they were present 
everywhere in swarms, and although at their worst in the forest, they were a pest 
in the villages, houses, trains, etc. Near the edge of the forest many more than a 
hundred Tabanids could be counted on one person at the same time, while hundreds 
of others would be flymg around. In the forest itself they were present in almost 
incredible numbers. There were many officers and men in the Force who had 
lived and travelled in all parts of the world, but all those I met agreed that never 
before had they encountered these flies in such swarms. 


196 MAJOR A. D. FRASER. 


Soon after the middle of July the flies diminished very considerably in numbers, 
and during the early part of August only an occasional fly or two would be seen 
on a particularly bright day. 

The flies seemed quite as eager to attack human beings as animals, and at their 
worst were a greater pest than I have ever found tsetse-flies. The majority of the 
flies settled on one’s legs, and in the forest they attacked in such numbers that work 
in it was rendered almost impossible. On one occasion I saw Lt.-Col. French, the 
Chief Liaison Officer, on his return from a reconnaissance through the forest ; the 
backs of his hands and his neck were completely discoloured, and his face was also 
badly discoloured as a result of bites. He said that as many as fifteen flies at a time 
would settle on the back of each hand, and his whole attention was occupied in 
attempting to ward off their attacks. 

The officers at Obozerskaya built a wooden tennis-court in the clearing, but the 
flies attacked so viciously and persistently, even in the evening between 6.0 and 
7.0 p.m., that the players were driven from the field, and compelled to give up the 
game until the flies diminished in numbers. 

Towards 10.0 p.m. the flies became less active, and between 10.30 p.m. and 
1.30 a.m., when only an occasional fly was to be seen, there was a welcome freedom 
from their attacks. At this season of the year in North Russia the sun only just 
dips below the horizon, and it is still quite light at midnight. 

The local inhabitants performed most of their outdoor work during the night, and 
kept their cattle and other animals in their sheds during the heat of the day. When 
a journey had to be made in the daytime, the drivers of the horses so far as possible 
selected certain times when they expected a wind would rise and cause the flies 
to become less troublesome. Before setting out they would rub their ponies with 
paraffin, a supply of which they would take with them. They would then proceed 
in convoys and make the journey as quickly as possible, using branches of trees to 
beat away the flies. The leading horse in the convoy seems to be attacked more 
severely than the others, and the animal’s head appears to be the favourite place of 
attack; but if blood has been made to ooze from any part of an animal this point 
always attracts a swarm of flies. As for human beings, I have heard it stated by British 
officers that they have been bitten through Bedford cord riding breeches. Everyone 
experienced much discomfort and annoyance from the constant attacks of these 
flies, and some individuals suffered from bites which developed into nasty septic 
sores requiring medical attention. 

_ It was noticed that a small, dark-coloured fly (probably a non-biting Muscid) 
constantly accompanied the Tabanids, and sucked up any blood that flowed from 
wounds caused by them. 
~ Tabanus.—The flies of this genus were the greatest pest, and always greatly 
outnumbered those belonging to Haematopota and Chrysops—at Obozerskaya, when 
Tabanids were at their worst, by something like 10 or 15 to 1, and 100 or 200 to 1, 
respectively. Tabanus No. 1 [T. tarandinus, L.—H.E.A.] was the first to appear, 
followed a day or two later by Tabanus Nos. 6 and 7 [T. maculcornis, Ztt., and 
T. mgrifacies, Gob.—E.E.A.]. The vast majority of the flies belonged to these 
three species until about the end of the first week in July, when they were gradually 
replaced by others. All these species were widely distributed throughout the 
district. 


NOTES ON BLOOD-SUCKING FLIES IN NORTH RUSSIA. 197 


Haematopota.—Representatives of this genus were found everywhere, and were 
most plentiful during the first three weeks in July. From ten to twelve were the 
most ever seen to settle on one person at the same time. Considering its numbers 
as compared with those of Tabanus, Haematopota appeared to be the more active 
biter. 

Chrysops.—First seen in the last week of June, Chrysops was met with until the 
end of the first week of August. Where the forest runs down to the edge of the 
Maliozerki Lake, as many as four or five of Chrysops No. 2 [C. caecutiens, L.— 
H.E.A.] would settle on one at once, but in the forest at some distance from water 
only one or two of these flies were encountered at a time. They were particularly 
fond of sitting on one’s cap. On cautiously removing my cap, after hearing the 
fairly characteristic buzz indicating the presence of one of these flies, I more often 
than not found the insect resting on it. 

Owing to their relatively small numbers, and their habit of rarely commencing 
to bite on first alighting, flies of the genus Chrysops were not a very great nuisance, 

Blood-Sucking Flies other than Culicidae and Tabanidae. Midges (Culicoides) 
and Simulium were troublesome everywhere. 

Phlebotomus was not found. 

Stomoxys were first seen at the end of June, but it was rather exceptional to come 
across more than an occasional specimen until August, when they could be found 
in fair numbers. 

Enemies of Blood-sucking Flies. Drayonflies, which were numerous throughout 
the summer, were frequently seen to capture specimens of the various species of 
Tabanus. 

Evidence of Disease. There was no evidence that any of the flies with which 
this paper is especially concerned conveyed any disease to human beings or domestic 
animals. 


APPENDIX. 
(By Major E. E. Austen, D.8.0.). 


The interesting statement by Major Fraser printed above confirms what is already 
known as to the extraordinary abundance and aggressiveness of TABANIDAE in 
Russia. Thus, according to the late Prof. Portschinsky (see Review Appl. Ent., 
ii, B, pp. 195-196), to whose long-continued researches previous knowledge of the 
subject is chiefly due, in Russia over 100 species of TABANIDAE are found, including 
some 60 species of Tabanus, 18 Chrysops, and 6 Haematopota, the remainder con- 
sisting of representatives of the genera Nemorius, Silvius, Pangonia and Heptatoma. 
In the polar regions of North Russia, where the number of species is limited, those 
which occur are frequently present in enormous numbers. According to an earlier 
paper by the same author (see Austen, British Blood-Sucking Flies, p. 33, 1906), 
in the Gdov District of the Petrograd Government, TABANIDAE in summer are so 
excessively numerous and bloodthirsty that agricultural operations have to be 
carried out by night ; while in parts of Siberia, such as the shores of the River Om, 
settlers have been compelled entirely to abandon the zone infested by these flies, 


198 MAJOR A. D. FRASER. 


The subjoined list, in addition to the names of the species represented in the 
collection brought back by Major Fraser and presented by him to the British 
Museum (Natural History), includes, for the sake of completeness, records derived 
from a study of one or two small series of specimens obtained in North Russia by 
other collectors during 1918-19, and recently added to the National Collection. 

Localities and other data are indicated as follows :—[F.]=Obozerskaya, between 
20. vi. and 31. viii. 1919 (Major Fraser). 

[F*.]= Chequevo, Onega River, end of June 1919 (Major Fraser). 

[B.]=250 miles up R. Dwina, vi. 1919 (Surg.-Lt. R. Buddle, O.B.E., R.N.). 

[C.]= Kola Gulf, vu. 1918, or other locality and date as stated (Capt. A. G. Carment, 

R.A.M.C.). Presented by Imperial Bureau of Entomology. 

[C*.]= precise locality and date unknown (Dr. E. A. Cockayne). 

* — determination by Mr. F. W. Edwards. 


CHIRONOMIDAE. 

Culicoides pulacaris, L. [F.] 

4s sp. incert.—Allied to C. festevipennis, Kieff. [F.] 

‘3 (2) arcuatus, Winn. [F.] 

»  fascrpennis, Staeg. . [F.] 
CULICIDAE. 

Anopheles maculipennis, Mg. hue 

Theobaldia siberrensis, Ludlow,* Archangel, 25. ix. 1918. [C.] 

Ochlerotatus alpinus, L.,* Murmansk, vi.-vii. 1918. [Cara 

fo lutescens, Fabr.,* Archangel, 1918. [C.] 
SIMULIIDAE. 

Simulium ornatum, Mg. OG bit baw 

, reptans, L. [C.] 

8 venustum, Say. [F.] 

; latipes, Mg. [Ct] 

zs sp. incert., near S. subexcisum, Edw. [C.] 

, hirtupes, Fries. [C.] 

TABANIDAE, 

Chrysops caecutiens, L. [F.] 

»  dwaricata, Lw. [F.] 

»  relicta, Mg. [F.] 

5  mgrupes, Ltt. [C.] 

,  sepuleralrs, Fabr. [Ft] 

Haematopota pluvialis, L. [F.][B.][C.] 

me crassicorms, Whibg. [F.} 

Tabanus tarandinus, L. [F. } 

»  troprcus, Pz. [F.] [B.] [C.] 

is s ,», var. bisignatus, Jaenn. [¥1} 

»  solstetialis, Schin. [PPB f 

,  luridus, Fin. [F.] [B.] 

5  negricornis, Ztt. [F.][F*] 

»  boreals, Mg. [F.] [C.]} 

,,  lapponicus, Whibg. [F.] 

»  maculicornes, Ztt. [F. ] 

»  mgrifacies, Gob. [F.] 
MUSCIDAE. 


Stomoxys calcitrans, L. [F.] 


199 


FURTHER NOTES ON THE LONCHAEIDAE (DIPT.), WITH DESCRIPTION 
OF NEW SPECIES FROM AFRICA AND ASIA. 
By fro, MW. Breet, 
Turin, Ltaly. 


Having received from the Imperial Bureau of Entomology more material of the 
family LoncHaErpAz, I am in a position to give the following additional notes to 
my previous paper* on the Ethiopian species. 

I think moreover it will be useful to offer a view of the Asiatic and Australian 
species at present known, as it seems that some of these flies (chiefly those of economic 
importance) are widely spread over the tropical countries of the Old World. Thus 
I have now found that the African plumosissima lives also in the Philippine Islands ; 
and that the Oriental citricola is to be found also in Australia. The wide range of 
aurea (splendida) is already known. 

THE SUBGENERA OF THE GENUS LONCHABRA, 8.L. 

In my previous paper (pp. 253-254) I have already shown that the genus 
Lonchaea may be divided into three natural groups, which are well characterised 
structurally and ethologically. These groups may be considered at present as 
subgenera, but further studies will probably show their generic rank. 

For the first group (type chorea, Fabr.) and the second group (type lasiophthalma, 
Macq.) there are already the names Lonchaea, s. str., and Dasyops, Rond. I will 
propose here the name Carpolonchaea, subgen. nov., for the mainly tropical species 
of the third group with feathered arista and with two sternopleural bristles (type 
plumossissima, Bezzi). The proposed name indicates the carpophilous habits of 
these species. 

A new subgenus is probably to be erected for the reception of the somewhat 
aberrant Lonchaea aurea (splendida), which has a rather isolated position in the 
genus Lonchaea, s. str.,on account of the different form of the head, the peculiar 
venation of the wings, the metallic colour of the body and the habits of the larva. 
The name Lamprolonchaea, subgen. nov., (type aurea=splendida) is proposed 
here for it. 

THe ETHiop1an SPECIES. 

In my previous paper 8 Ethiopian species of the genus Lonchaea were distinguished ; 
as I have now received 5 other species, thus bringing the total number to 13, the 
following new table of distinctions is necessary :— 

1(4). Antennae very short, widely separated at base, inserted a little below the 
middle of the eyes, with bare arista; eyes hairy ; head considerably 
broader than the thorax, with a very broad frons in the female, and with 
short and thin macrochaetae ; lunula very broad, open and roughly hairy ; 
cheeks and jowls broad ; sternopleurae with a single strong macrochaeta ; 
mesopleurae with anterior macrochaetae not developed ; wings shorter 
than the body, with the second costal cell not dilated outwardly, and with 
the sixth longitudinal vein destitute of spurious continuation (subgen. 
Dasyops, Rond.). 

*This Bulletin ix, 3, March 1919, pp. 241-254, figs. 1-4. 


8(11) 


(10). 


10(9). 
11(8). 


12(13) 


13(12). 


14(15). 


PROF. M. BEZZI. 


Cheeks smooth ; tarsi with the 3 or 4 terminal joints black ; base of wings 
and squamulae yellowish .. u At - .. mochu, Bezzi. 

Cheeks rugose ; only the last tarsal joint black; base of wings and squamulae 
blackish 4 ie oe a ape se phaeolepis, sp. n. 


Antennae longer, usually very long, closer together at base and inserted at 
or distinctly above the middle of the eyes ; eyes bare ; head not or only a 
little broader than the thorax, with less widened frons in the female, 
and with well developed macrochaetae ; lunula small, usually less visible 
and less or not hairy ; cheeks and jowls less broad, usually very narrow ; 
mesopleurae anteriorly with some strong macrochaetae, which are curved 
forwards ; wings longer than the body, with the second costal cell dilated 
outwardly and usually with the sixth vem with a long spurious con- 
tinuation. 


. Arista bare or only microscopically pubescent; only one sternopleural 


macrochaeta present ; tarsi always yellow at base ; squamulae whitish and 
with pale cilia. | 

Entire body, except head, of a very glistening green, more or less golden ; 
antennae short, with the third joint at most twice as long as the basal 
joints together ; jowls rather broad ; only the basal joint of all the tarsi 
yellow, with a black tip ; first posterior cell distinctly narrowed at end ; 
sixth longitudinal vein not at all continued (subgen. Lam~prolonchaea, 


nov.) 4) ve . aurea, Macq. 


Body of a shining bluish-black or black colour; 2 or 3 basal joints of tarsi 
yellow ; first posterior cell not narrowed at end; sixth longitudinal vein 
with spurious continuation (subgen. Lonchaea, s. str.). 

Antennae rather short, extending only a little below the middle of the face, 
wit the third joint at most twice as long as the basal joints together ; 
jowls rathe- broad. 

Vibrissae not longer than usual; wings yellowish hyaline . . . clar’pennie, 

Macq. 

Vibrissae long and strong ; wings brownish .. ihe vibrissifer, Lamb. 

Antennae long, extending to the epistome, with the third jomt many times 
longer than the basal joints together ; jowls linear. 

Tarsi with only the basal joint yellow ; third antennal joint extending below 
the mouth-border; wings brownish .. sik .. lambiana, Bezzi. 

Tarsi with 2 or 3 basal joints yellow; third antennal joint not extending 
below mouth-border ; wings hyaline, or faintly yellowish on fore half. 


Antennae quite black ; wings slightly yellowish on fore half 
continentalis, sp. n. 


NOTES ON THE LONCHAEIDAE (DIPT.) — 201 


15(14). Antennae with the interior part of 2nd and 3rd joint yellowish ; wings quite 
hyaline of .. wmpressifrons, sp. n. 

16(5). Arista plumose or at lease Seni iaaeodb ‘re Srond sternopleural macrochaetae 
present ; antennae always extending to the mouth-border, or even below, 
with the third joint many times longer than the basal joints together; jowls 
linear; tarsi entirely black ; first posterior cell not narrowed at end (subgen. 
Carpolonchaea, nov.). All the Ethiopian species here distinguished have 
white and pale hairy squamulae, and hyaline or pale yellowish wings. 

17(24). Plumosity of the arista of medium length, being not or only a little broader 
than the breadth of the third antennal joint ; the plumules are moreover 
thin, straight and close together. 

18(19). Last abdominal segment of the male longer than the two preceding ones 
together, deeply incised at end and there with the two points clothed 
with dense and long, bristly hairs... va excr-a, Kert. 

19(18). Last abdominal segment of the male not so eee | never specially ciliated. 

20(21). Abdomen at end with the two last ventral plates inflated and prominent ; 


smallish species with bare scutellum ae ..  gtbbosa, de Meij. 
21(20). Last ventral plates not so formed, never prominent; scutellum more 
hairy. 


22(23). Bluish species of smaller size (not over 4 mm.); plumosity of the arista 
broader than the breadth of third antennal joint ; wings quite hyaline 

laevis, sp. D. 

23(22). Black species of greater size, plumosity of the arista as broad as the third 

antennal,joint ; wings slightly but distinctly yellowish 

ophyroides, sp. n. 

24(17). Plumosity of the arista very long, being twice as broad as the breadth of 

the third antennal joint; the plumules moreover less thin, undulated 

and scattered i ie a ys plumosissima, Bezzi. 


1. Lonchaea (Dasyops) phaeolepis, sp. nov. 

Closely allied to mochiw, Bezzi, from Erythraea, but smaller and at once 
distinguished by the characters given in the table. 

3S 9. Length of the body, 3-32 mm.; of the wing, 2°9-3 mm. 

Head and its appendages as described for mochiw, with the following differences : 
the frons lacks the greenish reflexions ; the cheeks in the upper part and the sides of 
frons in the anterior part with rather developed transverse furrows, being thus 
strongly rugose ; arista yellowish at base. Thorax and scutellum without distinct 
aeneous reflexions, and with the same chaetotaxy. Squamulae blackish, and with 
a blackish fringed border ; halteres black, with paler stalk. Abdomen as in mochii, 
but the ovipositor is distinctly shorter and broader. Legs of all the pairs with the 
tarsi entirely yellow, the last joint only being black. Wings proportionally shorter 
and broader, with the base deep blackish to the basal cross-veins; the first 
posterior cell is not distinctly narrowed at end; the sixth longitudinal vein is shorter, 

Type ¢ and type 9, anda single couple of specimens (British Museum) from 
Durban, Umbilo, 16.xi.1913 and 23.11.1914 (L. Bevis). In the Cape Museum 
there is a specimen from Natal. 


202 PROF. M. BEZZI. 


2. Lonchaea (Lamprolonchaea) aurea, Macq. 

Of this interesting species I have seen in the Museum of Genoa specimens taken 
at S. Nicolas and 8. Iago, Cape Verde Islands, in May and November 1898, by the 
late L. Fea ; and in the Cape Museum specimens from the Transvaal. 


3. Lonchaea (Lonchaea) continentalis, sp. nov. 

Very closely allied to lambiana, Bezzi (longicornis, Lamb), from the Seychelles, 
and perhaps only a continental form of it, but seeming to differ in having the tarsi 
more broadly yellow at base, and in the wings not being brownish. 

gd. Length of the body, 3-32 mm. ; of the wing, 3°1-3°3 mm. 

Head and its appendages entirely black ; frons proportionally narrow, opaque, 
with shining vertical plates ; lunula below white, shining ; face quite flat and clothed 
with whitish dust, which is more shining at sides ; antennae with the third joint 
extended to the mouth-border and with long, black, microscopically pubescent 
arista ; palpi broad, with some bristly hairs. Chaetotaxy normal. Thorax and 
scutellum shining black, not dusted at all, with bluish reflexions on the back; chaeto- 
taxy normal; mesopleurae with bristly hairs on the middle, 2-3 bristles curved 
backwards at hind border, and 2 bristles curved forwards at anterior border ; 
sternopleurae with only one strong bristle ; scutellum with some hairs at border 
between the usual bristles. Squamulae white and white-fringed ; halteres black. 
Abdomen shining black and black-haired ; it has no special ciliation at the tip, nor 
prominent genitalia ; the last segment is simple and only a little longer than the 
preceding one. Legs black, with the two basal joints of all the tarsi yellow. Wings 
hyaline, slightly greyish on the fore half ; second longitudinal vein quite straight ; 
terminal portions of the 3rd and 4th veins parallel ; small cross-vein a, little before 
the middle of the discoidal cell ; the spurious continuation of the 6th vein is only 
faintly indicated ; second costal cell with normal dilatation. 

Type ¢ and an additional specimen of the same sex (British Museum) from Natal, 
Durban, 28.ix. 1916 (C. P. v.d. Merwe): another male specimen in the writer’s 
collection from Erythraea, Ghinda, vin. 1916 (Dr. A. Mocha). 


4. Lonchaea (Lonchaea) impressifrons, sp. nov. 

Very similar to the preceding species, but of a more glistening bluish colour, with 
partly reddish antennae and with crystalline wings. 

©. Length of the body, 2°6 mm. ; of the wing, 2°8 mm. 

Frons rather narrow for a female, opaque, with shining ocellar and vertical plates ; 
before the ocelli there is a distinct ovate depression ; lunula shining white; face 
rather shining, with a faint greyish dust. Antennae extending to the mouth-border, 
with the second joint and the rather broad third joint broadly reddish on the interior 
side, the last only on the basal half; arista not very long, miscroscopically pubescent ; 
palpi and proboscis black. Thorax and scutellum very glistening bluish ; chaeto- 
taxy as In the preceding species, scutellum with 2-3 bristly hairs. Squamulae and 
halteres as in the preceding. Abdomen very shining, but less bluish than the back of 
mesonotum ; ovipositor about 1 mm. long. Legs black, the tarsi with the two basal 
joints and part of the third joint pale yellowish. Wings as in the preceding species, 
but proportionally broader, they are quite crystalline, even at fore border ; veins 
very pale yellowish. 


NOTES ON THE LONCHAEIDAE (DIPT.). 203 


Type 9, a single specimen (British Museum) from Gold Coast, Accra, iv. 1916, 
“in laboratory ” (Dr. J. W. S. Macfie). 


5. Lonchaea (Carpolonchaea) laevis, sp. nov. 

A shining, bluish-black species with quite hyaline wings, distinct in the male sex 
owing to the not prominent genitalia and the simple last abdominal segment ; from 
plumosissima it is distinguished by its smaller size and the shorter plumosity of the 
arista. 

$ 9. Length of the body, 3°5-3°8 mm. ; of the wing, 3°8—4 mm. 

Frons black, opaque or with faint sericeous reflexions, the vertical and ocellar plates 
bluish and shining; in the female it is only 1} times as broad as that of the male. 
Antennae entirely black, with the third joint extended to the epistome, three times 
as long as the basal joints together ; arista yellowish at base, its plumosity broader 
than the breadth of the third antennal joint, the plumules close together and straight. 
Lunula black, but the prominence between the base of the antennae is white-dusted ; 
face greyish-dusted ; palpi and proboscis black ; jowls linear, vibrissae not specially 
developed. Thorax very glistening on the back and there with bluish reflexions, 
black on the pleurae ; mesopleurae on anterior border with 3, and on posterior border 
with 3 bristles, in the middle with a tuft of bristly hairs ; sternopleurae with 2 strong 
bristles, which are curved upwards. Scutellum more aeneous, faintly dusted, with 
short bristly hairs at border. Halteres black ; squamulae white and with white 
cilia. Abdomen coloured and shining like the back of mesonotum but without 
bluish reflexions ; last segment of male not elongated or ciliate ; genitalia not visible ; 
ovipositor as long as the last two segments together. Legs entirely black ; middle 
femora with complete and long ciliation on hind side. Wings quite hyaline and with 
pale yellowish veins ; second costal cell very much dilated outwardly ; last portions 
of 3rd and 4th veins parallel ; small cross-vein before the middle of the discoidal cell 
and before the end of the first longitudinal vein ; chitinised part of the sixth vein 
short, but the spurious continuation extended to the hind border ; hind cross-vein 
quite straight, and longer than its distance from the end of the fifth vein. 

Type 3, type 2 and an additional specimen (British Museum) from Zanzibar (Dr. 
W.M. Aders) ; some other specimens of both sexes in the writer’s collection from 
Hrythraea, Ghinda, August-December 1916 (Dr. A. Mochi). 


6. Lonchaea (Carpolonchaea) ophyroides, sp. nov. 

Also closely allied to plumosissima, but easily distinguishable on account of its 
greater size and the much shorter plumosity of the arista. 

3 9. Lengths of the body, 4°2—4'5 mm. ; of the wing, 455—4°8 mm. 

Head as described for plumosissema, but the antennae with the third joint not 
extending below the oral margin, and with the arista much more shortly plumose, 
the breadth of the plumosity being not broader than the breadth of the third antennal 
joint and the plumules being thin, dense, and quite straight. All the rest of the body, 
legs and wings as in plumosissima ; but the abdomen is aeneous, without bluish 
reflexions ; the ciliation at hind side of middle femora is more developed ; the wings 
are distinctly yellowish-fuscous on the fore half. 


204 PROF. M. BEZZI., 


Type 3, type 2 and an additional female specimen (British Museum) from British 
Hast Africa, Kabete, 28.vii.1914 “on window” (7. J. Anderson); a female 
specimen without antennae from N. Nigeria, Zarda, “ ovipositing in vegetable 
marrow ” (P. H. Lamb) seems also to belong here. 


7. Lonchaea (Carpolonchaea) plumosissima, Bezzi. 

In fully coloured specimens the wings are sometimes yellowish fuscous on the 
fore half. ? 

There are specimens from the type locality, Gold Coast, Aburi, January—February 
1911, “from kola pod” (L. Armstrong); and from the same locality, 1912-13 
(W. H. Patterson) ; and moreover a male specimen from British East Africa, Embu, 
24.vii.1914 (G. St. J. Orde-Browne) ; thus showing that the species is present even 
in the east of the Ethiopian Region. 

Puparvum. Of the specimens reared from kola pods there is also a puparium, 
this is elongate, about 4 mm. long, of a shining reddish colour ; the segmentation 
is not well marked; the skin is hard, finely rugulose ; the posterior spiracular 
tubercles are small, black, rounded, approximated, and projecting for about the 
length of their diameter. 


THE Inpo-AUSTRALIAN SPECIES. 

The Indo-Australian species of Lonchaea, s.l., were tabulated by Dr. Kertész 
as long ago as 1901; many species have been added since by Prof. de Meijere and 
by me. All the subgenera are well represented, with the exception of the subgenus 
Dasyops, of which no Oriental or Australian species are at present known; even 
in the very numerous Dipterous galls from Java, described by W. and J. Docters 
van Leeuwen-Reijnvaan, 1901-1916, it seems that there are no galls referable to 
these flies. 

The species at present known, with the addition of some new ones, can be distin- 
guished as follows :— 

1(8). Arista bare or only microscopically pubescent ; only one sternopleural bristle ; 
legs always with the tarsi partly yellow; squamulae white or pale 
yellowish and with whitish cilia.* 

2(3). Metallic species of golden green colour, with rather short antennae and 
rather broad jowls; first posterior cell distinctly narrowed at end ; 
tarsi with only the basal joint yellow, with a black tip . . . aurea, Macq. 

3(2). Black or bluish-black species with narrow jowls and long antennae, the third 
joint of which is many times longer than the basal ones and reaches the 
epistome ; first posterior cell not narrowed at end; tarsi more broadly 


yellow. 
4(5). Only the basal joint of all the tarsi yellow, but that entirely so; wings 
yellowish ; species more than 4 mm. long... ..  megacera, Kert. 


5(4). The second joint of all the tarsi also yellow; wings wholly hyaline; species 
not over 3 mm. long. 


* The Lonchea ? consentanea, Walker 1860, from Macassar and Gilolo, of which the 
author says nothing about the colour of tarsi, is described as having white halteres, and 
thus cannot be a member of this genus. 


NOTES ON THE LONCHAEIDAE (DIPT.). 205 


6(7). Hypopygium very small, without pen mar APeRnEAers thorax black ; 


jowls rather broad . oe minuta, de Meij. 
7(6). Hypopygium with a ane Sirved appendage aa ‘thorax aeneous ; 
jowls linear .. ; re pugrnata, de Meij. 


8(1). Arista more or less eee ans even if sometimes rather shortly ; two sterno- 
pleural bristles ; legs with entirely black tarsi (with the single exception 
of pallicarpa) ; jowls always narrow; antennae always long, with the 
third joint many times longer than the basal joints together and reaching 
the epistome.* 

9(20). Squamulae dark or blackish, and with blackish cilia ; thorax and scutellum 
never pollinose. 

10(13). Wings hyaline and with pale yellowish veins. 

11(12). Arista with long plumosity ; scutellum with numerous bristly hairs at 
border between the usual bristles ; length of body about 5 mm. 

citricola, Bezzi, 3. 

12(11). Arista with shorter plumosity ; scutellum with a few short hairs near the 
apex; length of body about3mm. .. ed ficuperda, Bezzi, 3. 

13 10). Wings very dark, blackish or brownish, with dark veins. 

14(15). Third antennal joint a little yellowish near the base ; hypopygium with a 
broad and strong appendage below; smaller species, about 3 mm. long. 

cupraria, de Mei. 3. 
15(14). Third antennal joint quite black; hypopygium not so formed, without 
| prominent appendage below, species 4-5 mm. long. 


16(17). Abdomen violaceous, shining. . A 4 iM ..  biror, Kert. 3. 

17(16). Abdomen black. 

18(19). Abdomen rather opaque in the middle a r. atratula, Walk. g.t 

19(18). Abdomen entirely shining .. i is obscurupennis, de Meij. 3. 

20(9). Squamulae white or pale yellowish, al whitish cilia. 

21(22). Tarsi yellowish ; scutellum bare ; wings broad, with the third longitudinal 
vein ending before the tip of wing a ahs pallicarpa, sp. nov. 

22(21). Tarsi entirely black ; wings less broad, with the third vein ending at tip of 
wing. 


23(26). Arista rather shortly plumose, the breadth of the plumosity being narrower 
than the breadth of the third antennal joint; scutellum not or less 


pilose. 
24(25). Thorax and scutellum not pollinose ; 3 mm. ae ..  calva, Bezzi. 
25(24). Thorax posteriorly and scutellum pollinose ; 4 mm. .. pollinosa, Kert, 


26(23). Arista with longer plumosity, which is at least as broad as the third joint, 
27(28). Abdomen dull black, a little shining towards the base of the sides ; 3 mm. 
montana, Brun. 
* The species Lonchaea ? punctipennis, Walker 1860, from Macassar, and Lonchaea? inops, 
Walker 1859, from Aru, Mysol and Ceram, evidently do not belong to the present genus. 
+Walker, who describes only the female, says nothing about the colour of the squamulae, 
which are assumed here to be blackish (according to the colour of the wings), at least in 
the male sex. 
(713) B 


206 PROF. M. BEZZI. 


28(27). Abdomen entirely shining. 
29(38). End of the abdomen in the male without special conformation and without 
prominent genitalia.* 
30(37). Plumosity of the arista not twice as broad as the third antennal joint, with 
straight plumules. 
31(32). Wings brownish, darker on the fore border and apical half. 
albisquama, Kert. 


32(31). Wings hyaline or more or less deep yellowish. 

33(36). Species of greater size (4-5 mm.), with deep yellow wings. 

34(35). Frons of the female with 3 longitudinal furrows .. cyaneomtens, Kert. 

35(34). Frons of the female with 2 furrows only... 56 bisulcata, sp. nov. 

36(33). Species of smaller size (3°5 mm.), with hyaline or slightly yellowish wings ; 
frons of the female not distinctly sulcate 2 jilifera, Bezzi. 

37(30). Plumosity of the arista twice as broad as the third jomt, with undulate 
scattered plumules .. se ot is .“ plumosissima, Bezzi. 

38(29). Abdomen of the male at end with special conformation or with prominent 
genitalia. 


39(40). Last abdominal segment of the male twice as long as the preceding one, 
deeply excised at end and there with the 2 points clothed with long bristly 
hairs; hypopygium small ap : ei excisa, Kert. 

40(39). Last abdominal segment not so elongate sinc not pacts at end. 

41(42). Last abdominal segment simple, but with long black hairs at sides, ciliated ; 
hypopygium small and not promiment below, but with a horizontally 
produced middle appendage ae ae setifera, de Meij. 

42(41). Last abdominal segment not specially ce at sides; hypopygium very 
prominent below. 

43(44). Hypopygium properly small, but the last two ventral segments inflated 
and prominent in the shape of a rounded protuberance. 

gibbosa, de Meij. 

44(43). Abdomen at end not gibbous beneath, but the genitalia prominent in the 

shape of a broad and stout appendage, which is bilobate at end 
lucens, de Meij. 


1. Lonchaea (Lamprolonchaea) aurea, Macq. 1851. 
Lonchaea splendida, Loew, 1873. 
Lonchaea metatarsata, Kertész, Term. Fiizet., xxiv, 1901, p. 83; de Meijere, 
Tijds.° v, Ent., li, 1908, p.°133 ; lin, 1910, p. 117; Ivui, 1915, 80; 
lx, 1918, p. 350; Edwards, Trans. Zool. Soc., xx, 1915, p. 422. 

This little fly seems to be the most widely spread species of its genus; in the 
Orient it is recorded from South India. The synonymy with metatarsata is quite certain, 
as I have seen a specimen from Batavia (Jacobson) in the Indian Museum, deter- 
mined by Prof. de Meijere. JL. metatarsata was originally described from New 


* Of cyameonitens and bisulcata only the females are known ; the males are here assumed 
to have a simple abdomen. 


NOTES ON THE LONCHAEIDAK (DIPT.). 207 


Guinea, and Prof. de Meijere has recorded it from Java and Sumatra. L. splendida 
is recorded from Australia, New South Wales ; and Froggatt (Austr. Insects, 1908, 
p. 308) says that it has a wide range from the Pacific Islands and New Zealand, over 
Australia ; thus the L. splendida, Broun 1905, for which I have proposed the new 
name of browmana, is probably the same species, 

[ have before me specimens from the Philippine Islands: Luzon, Laguna, Los 
Bafios and Mt. Maquiling (C. F. Baker;) and Panay Culasi, vi. 1918 (R. C. 
MacGregor). 


2. Lonchaea (Lonchaea) minuta, de Meijere. 
Lonchaea minuta, de Meijere, Tijds, v. Ent., liii, 1910, p. 116, and lx, 1918, p. 350. 


Described from Java, I have before me a specimen from Malacca, Kelantan, and 
another from the Philippines, Batbatan Isl., vi. 1918 (R. C. MacGregor). 


3. Lonchaea (Carpoionchaea) citricola, Bezzi. 

Lonchaea citricola, Bezzi, Philipp. Jl. Sci., vii, D., 1913, p. 319. 

Of this species, described from the Philippimes, Los Bafios,there are in the collection 
submitted to me by the Imperial Bureau two male specimens from Australia, N. 
Territories, Darwin, 1l.v.1914 (G. F. Hull), They agree perfectly with the types, 
and have been bred from larvae from rotten oranges. 


4, Lonchaea (Carpolonchaea) ficiperda, Bezzi. 
Lonchaea ficiperda, Bezzi, Philipp, Jl. Sci., vii, D., 1913, p. 320. 


I have seen other specimens of this species, likewise from the Philippines, 
Batbatan Isl., vi. 1918 (R. C. Mactrregor). 


5. Lonchaea (Carpolonchaea) atratula, Walker. 

Lonchaea atratula, Walker, Proc. Linn. Soc. Zool., iv, 1860, p. 146; Kertész, Term. 
Fiizet., xxiv, 1901, p. 84. 

If I have correctly interpreted this species, it is distinguished by its greater size, 
blackish and dark-fringed squamulae, blackish wings and the dullish disc of the 
abdomen. It is probable that birov, Kert. from New Guinea, and obscuripennis, 
de Meij., from Java, belong to this same species. 

Originally described from Macassar, I have seen specimens from Singapore (C. F. 
Baker) and from the Philippines, Mindanao, Butuan (C. F. Baker). 


6. Lonchaea (Carpolonchaea) pallicarpa, sp. nov. 

Very distinct from all the other known species of the subgenus Carpolonchaea from 
the Old World on account of the yellowish tarsi; characteristic are also the broad 
wings, with diverging third and fourth veins. 

Q. Length of the body, 3 mm. ; of the wing, 3:2 mm. 

Head black ; frons flat, not sulcate, broad, being only a little longer than broad, 
with sericeous reflexions on the middle, and with glistening and bluish vertical plates. 
Antennae black, with the third joint a little brownish along the inner border and 
reaching the epistoma, being about three times as long as the basal joints; arista 
with long plumosity which is as broad as the breadth of the third antennal joint ; 

(713) B2 


208 PROF. M. BEZZI. 


the plumules thin and straight. Vibrissae and lateral bristles of the mouth-border 
but little developed ; palpi and proboscis black. Thorax black and very shining, 
with distinct bluish reflexions on the back; dorsal hairs very short; chaetotaxy 
normal; mesopleurae with two anter or and three posterior bristles; two strong 
sternopleural bristles. Scutellum rather aeneous and faintly dusted, quite bare 
except for the usual bristles ; squamuiae white and white-fringed ; halteres black. 
Abdomen like the mesonotum, but without bluish reflexions; ovipositor 1 mm. 
long. Legs black, with the two basal joints of all the tarsi reddish yellow ; middle 
femora with complete ciliation on hind border. Wings proportionally broad, quite 
hyaline, with pale yellowish veins ; the terminal portions of the 2nd, 3rd and 4th 
veins are straight, gradually and equally diverging, so that the third ends a little 
before the tip of the wing ; small cross-vein before the middle of the discoidal cell ; 
hind cross-vein straight, only a little longer than its distance from the end of the 
fifth vein, sixth vein with spurious continuation to the hind border. 

Type 9, a single specimen in Prof. Baker’s collection from the Philippines, Baguio, 
Benguet (C. F. Baker). 


7. Lonchaea (Carpolonchaea) bisulcata, sp. nov. 

A shining black species of proportionally greater size, with deep yellow wings; 
distinguished by the bisulcate frons of the female. 

Q. Length of the body, 4 mm. ; of the wing, 4:2 mm. 

Frons proportionally narrow, a little less than twice as long as broad ; it is dullish 
black, with sericeous reflexions and with shining black vertical plates; in front of 
the ocellar plate there is a rounded depression, from the sides of which emerge 
two rather deep furrows, which converge towards the antennae, ending separately 
at the upper border of the lunula. Antennae wanting in the type. Face black, 
greyish-dusted, with rather shining antennal grooves ; cheeks linear ; jowls narrow ; 
palpi and proboscis black ; chaetotaxy normal. Thorax shining black, with no dis- 
tinct bluish reflexions ; chaetotaxy normal ; two equally strong sternopleural bristles. 
Scutellum aeneous and faintly dusted, with numerous bristly hairs between the 
usual bristles. Squamulae yellow, with pale yellowish cilia; halteres black. 
Abdomen coloured and shining like the mesonotum, with black hairs and short 
black bristles on the sides; ventral membrane broad and reddish, ventral plates 
shining black ; ovipositor short, as long as the last two abdominal segments together. 
Legs entirely black; middle femora with rather short but complete ciliation on 
hind side. Wings yellowish, deeper yellow at base and on fore half ; veins yellowish. 
but the costa appearing darker on account of its short black ciliation ; second costal 
cell but little widened outwardly ; terminal portions of third and fourth veins parallel, 
not diverging ; small cross-vein before the middle of the discoidal cell ; hind cross- 
vein straight, about twice as long as its distance from the end of the fifth vein ; sixth 
vein with spurious continuation. 

Type 9, a single specimen in the writer’s collection from South India, Trichinopoly, 
1911 (fF. Cajus); the chaetotaxy of the thorax and the black tarsi, in spite of the 
want of the antennae, show that the species belongs to the subgenus Carpolonchaea. 

The present species may be the Indian Lonchaea without specific name figured by 
Howlett in Maxwell-Lefroy, Indian Insect Life, 1909, p. 636, fig. 420, from Pusa, 


NOTES ON THE LONCHAEIDAE (DIPT.). 209 


of which he says, “ fairly common blue-black Lonchaea sometimes seen sitting on 
leaves in the neighbourhood of excrement.”’ But these last words may prove that 
it is a Lonchaea, s. str. 


8. Lonchaea (Carpolonchaea) filifera, Bezzi. 

Lonchaea filifera, Bezzi, Philipp. Jl. Sei., vii, D., 1913, p. 320. 

Of this species, which seems to be common in the Philippine Islands, I have seen 
other specimens from Luzon, Laguna, Los Bafios and Mt. Maquiling (C. F. Baker), 
and from Manila, x. 1918 (R. C. MacGregor). 

The yellow penis is not always visible ; the female is characterised by the broad, 
dull black frons. 


9. Lenchaea (Carpolonchaea) plumosissima, Bezzi, 1919. 

Of this African species I have seen a male specimen from the Philippmes, Luzon, 
Los Bantos (C. F. Baker), which agrees perfectly in the arista with the typical 
specimens from the Gold Coast ; it has a deep black, quite opaque frons, and the 
wings deep yellowish-fuscous on the fore half. 


10. Lonchaea (Carpolonchaea) excisa, Kertész. 


Lonchaea excisa, Kertész, Term. Fiizet., xxiv, 1901, p. 87; de Meijere, Tijds. 
v. Ent., li. 1908, p. 133 ; lin, 1910, p. 118, pl. 7, fig. 50 ; Ix, 1918, p. 350. 

Of this, in the male sex, very characteristic species, originally described from 
Singapore and recorded from Java and Stmalur, there are in the Imperial Bureau 
Collection specimens from Ceylon, Peradeniya, 1918 (N. K. Jardine), and from the 
same locality,. 8.vi:1915 (Green) ; also from South India, Coimbatore, 17.vui.1912. 
In the Philippines it is a common species, caught in different localities of Luzon 
and of Mindanao by C. #, Baker and R. C. MacGregor ; I have also seen specimens 
taken at Singapore by Prof. Baker. 


11. Lonchaea (Carpolonchaea) setifera, de Meijere. 
Lonchaea setrfera, de Meijere, Tijds. v. Ent., liii, 1910, p. 119, pl. vu, fig. 52; lx, 
1918, p. 350. 

Even this species, described from Java and recorded from Simalur and New 
Guinea, is common in the Philippines, as I have seen specimens from Los Bafios 
and Mt. Maquiling (C. F. Baker); from Manila (R. C. MacGregor) ; from Leyte, 
Tacloban (C. F. Baker.) 


12. Lonchaea (Carpolonchaea) lucens, de Meijere. 
Lonchaea lucens, de Meijere, Tijd. v. Entom., liii, 1910, p. 118, pl. vii, fig. 51, 
lx, 1918, p. 350. 
Originally described from Java and recorded from Sumatra, I have seen specimens 
from the Philippines: Luzon, Los Bafios (C. F. Baker) and Panay, Culasi, v. 1918 
(Rh. C. MacGregor). 


210 PROF. M. BEZZI. 


ON THE FEMALES OF THE SUBGENUS CARPOLONCHAEA FROM THE ORIENT. 


The females of this group are for the most part unknown, even Prof. de Meijere 
having described only males. Those known to me are not easy to distinguish. 
I have not seen females of the species with dark squamulae, except that of my 
fictperda, which has however whitish squamulae ; it 1s therefore possible that even 
the species citricola and atratula have the squamulae differently coloured in the two 
sexes. Of birdi,cupraria and obscuripennis only the males are known ; of atratula, 
Walker, the author has described the female, but it is possible that he was mistaken 
as to the sex of his specimen. 

The females known to me may be distinguished as follows : 

1(2). Arista with very short plumosity, which is much narrower than the breadth 
of the third antennal joint 83 .. calva, Bezzi. 
2(1). Plumosity of the arista longer, at least bial to the breadth of the third 
antennal joint, or nearly so. 
3(4). Tarsi yellowish ; frons broad ; third and fourth longitudinal veins diverging 
| pallicarpa, Bezzi. 
4(3). Tarsi quite black ; third and fourth veins parallel at end. 
5(16). Plumosity of the arista not twice the breadth of the third antennal joint, 
with straight plumules. 
6(11). Frons broad, not twice as long as broad. 


7(8.) Frons of an opaque, deep black colour ; arista rather shortly plumose 
jilrfera, Bezzi. 


8(7). Frons rather shining, with sericeous reflexions ; arista with longer plumosity. 
9(10). Wings yellowish fuscous .. att . a .. lucens, de Mei. 
10(9). Wings hyaline bie re a ..  setifera, de Meij. 
11(6). rons narrower, twice as iar as iaoual 
12(13). Ovipositor about as long as the abdomen .. “% .. jficiperda, Bezzi. 
13(12). Ovipositor much shorter than the abdomen. 
14(15). Wings deep yellowish ; frons with a deep depression and with two longitu- 


dinal furrows converging anteriorly ; species of greater size 
bisulcata, Bezzi. 
15(14). Wings hyaline or slightly yellowish ; frons without basal depression and 
without marked furrows ; species of smaller size .. excisa, Kert. 
16(5). Plumosity of the arista twice as long as the breadth of the third antennal 
jot, with undulated plumules, which are more thick and more scattered 
than usual; frons narrow oe i .. plumosissima, Bezzi. 


211 


ON THE BRITISH SPECIES OF SIMULIUM.—II. THE EARLY STAGES ; 
WITH CORRECTIONS AND ADDITIONS TO PART I. 
By F. W. Epwarps. 
(Published by permission of the Trustees of the British Museum.) 


In the introduction to the first part of this paper* it was remarked that very little 
exact work had been published on the adult European Simutmpar. This is 
equally true of the larvae and pupae; in fact no one has attempted to describe the 
early stages of more than two or three of the common species. Moreover, in spite 
of their great interest, and the ease with which they can be collected and reared, 
they have apparently been almost entirely ignored by collectors, so that whereas in 
publishing my previous work I was able to acknowledge valuable assistance from a 
number of fellow entomologists, the results given in the following pages are, unless 
otherwise stated, due to my own observations. I wish especially to thank Mr. M. E. 
Mosely for the early stages of S. subornatum, the last species to come under my notice 
in the pupal stage. While there are still many important gaps in our knowledge 
which remain to be filled, it has been thought worth while to state without further 
delay the results that have been obtained so far. 

The purpose of the present paper is to give the writer’s observations on the biology 
of the insects, and to characterise, sufficiently for purposes of identification, the 
known larvae and pupae. In the course of rearing some species from the pupae, 
it was discovered that the females of the latipes group had been associated previously 
with the wrong males, and their characters had been badly confused, both im my 
original paper and in the reprint of it whichappeared in the Entomologists’ Monthly 
Magazine. Owing to this somewhat serious error, which has led the writer to adopt 
a different interpretation of Fries’ S. awreum, and owing also to the discovery of 
four apparently new species, it has been thought advisable to tabulate afresh the 
adults of both sexes. I am indebted to Mons. E. Ségny,of the Paris Museum, for 
examining at my request Meigen’s types of Simulium ; he reports that in ornatum, 
replans, argyreatum, sericatum and latvpes of Meigen, the types agree with the 
interpretation here given. 

From the systematists’ point of view the most valuable memoirs which have 
appeared dealing with the early stages of Simulivum are those of Lutz}, Mallocht 
and Pomeroy § on the American species. To all these writers the present author is 
glad to express his indebtedness, As regards the early stages of the SIMULIIDAE 
outside Britain and North and South America, they are still scarcely known, though 
Mr. Pomeroy has lately undertaken the study of the African species. | 

* Bull. Ent. Res., vi, pp. 23—42 (1915). 


+ Lutz, A., Mem. Inst. Oswaldo Cruz, i, p. 124 (1909), and ii, p. 213 (1910). 

~ Mattocn, J. R. ‘American Black Flies or Buffalo Gnats.’-—U.S. Dept. Agric. 
Bur. of Ent. Tech. Series 26, 1914. 

§ Pomeroy, A. W. J. ‘‘ Notes on five North American Buffalo Gnats of the genus 
Simulium.”—U.8. Dept. Agric. Bull. 329, 1916. 

| While this work has been passing through the press a valuable paper on the German 
species has been published by Dr. K. Friepricus (Vorlauf. Mitt. Sitzb. Abh. Naturf. 
Ges. Rostock, vii, pp. 211-226, Oct. 1920). The adults, pupae and habitats of nine 
species are briefly described). 


212 F. W. EDWARDS. 


Before passing to tabulate the species and consider them individually there are a 
few points which seem to be of sufficient general interest to claim attention here. 


Method of Rearing and of Association of Larvae and Pupae with Adults.It was pointed 
out by Newstead* that the pupae of Semuliwm when removed from the water and 
placed in a moderately damp atmosphere without superfluous moisture will readily 
complete their transformation into adults. This method of rearmg has proved 
invaluable, and provided the vessel in which the pupae are kept is not too small, and 
is kept slightly moist, it practically never fails : thousands of specimens have been 
reared in this way. If however a male pupa which it is desired to rear should die, 
it can easily be identified by dissecting out the genitalia. 

In a colony of larvae where even a moderate proportion are full-grown, some are 
sure to be found which exhibit a round black spot on each side of the prothorax, 
conspicuous even to the naked eye. This is the respiratory organ of the developing 
pupa showing through the larval skin, its component filaments bemg coiled up into 
a small space. The filaments can be very easily dissected out with a pair of needles, 
and a positive identification of larva and pupa obtained. If in the same colony 
there are also blackish, well-developed pupae, the species can be identified by dis- 
secting these without the necessity of waiting for specimens to hatch out. It is very 
desirable to identify larvae by this reliable method, since one very frequently finds 
several species living together in the same colony; though in a small area of a few 
square inches one will usually greatly outnumber the others. 


Over-wintering.—Some species, notably S. ornatum, appear to contimue breeding 
throughout the winter, even in the coldest weather, though, as might be expected, 
the rate of development is greatly retarded. Of such species there must be at least 
three broods in the year. Others, such as S. latvpes, pass the writer as young larvae, 
developing rapidly in the early spring ; probably these are normally single-brooded. 
The species which appear in such vast numbers in spring in our muddy-bottomed 
rivers may possibly winter in the mud in the egg stage, but I have not been able to 
obtain any positive evidence of this. Another possibility,and perhaps a more likely 
one, is that the females may hibernate, but 1 have never found them in a torpid 
state. However, as Mr. Hamm has pointed out to me, the specimens of S. equinum. 
and S. argyreatum captured very early in the year are generally, if not always 
females. More observations are needed on this point. 

Oviposition.—V ery little is yet known as to the egg-laying habits of the different 
species, but it would seem that great variation exists. I have been able to confirm 
the very interesting observations of Mr. H. Britten} on S. equinum, and have added 
some further particulars below. J have also found and hatched the eggs of S. aureum, 
which, unlike those of S. equinum, are hard-shelled and not enclosed in a mass of jelly; 
the newly-hatched larva of this species also has a much more strongly-developed 
egg-burster, so that it would seem that this species may be adapted to resist 
desiccation in the egg state. 

Habitat of Larvae.—The larvae which I have met with may be classified according 
to their habitat into (1) those which live on water-plants (or other objects if available) 


* Ann. Trop. Med., i, p. 40. 
{ Ent. Mo. Mag. (3), i, 1915, p. 150. 


ON THE BRITISH SPECIES OF SIMULIUM—II, 213 


in sluggish and permanent rivers such as those of East Anglia ; (2) those which live 
on rocks, stones (chiefly the upper sides) and sometimes on water-plants in swift 
mountain streams, which may become dry in dry seasons ; and (3) those which live 
chiefly on the undersides of stones, in small temporary streams, which are dry for 
several months of the year. These classes are not rigidly defined, ether in regard 
to the water-courses or to the insects, but it is often possible to say from the 
appearance of a stream what species will be found in it. Some, for example S, latipes, 
are more particular in their choice of a locality than others, such as S. ornatum, 
which is perhaps the most catholic in its tastes. The different species found in one 
type of stream are noi all closely related, as is shown by a study of the adults and 
pupae ; but it is a noteworthy fact that most of the larvae which live in very rapid 
streams have their anal gills very much branched, while most of those which live in 
sluggish rivers have the three mai branches simple. It might have been supposed 
that the more rapidly-moving and presumably more highly aerated water would not 
need such a large surface in the gills for the insects to accomplish their respiration, 
but this is not what we actually find. It may be that the presence of water-plants 
assists respiration to an even greater extent than movement of the water. 

Larvae always occur in the greatest numbers in the swiftest part of whatever 
stream they are in. The slow-river species furthermore show a decided preference 
for those water-plants (e.g., Scorpus, Sparganium and Ranunculus) which have long 
and narrow or much divided leaves, the reason perhaps being that such leaves are 
waved about by the current and thus provide better aeration ; the larvae occur 
most abundantly towards the tips of the leaves. 

The species found on stones all show a decided preference for clean stones ; 
when there is much moss or algal growth orgccumulation of sediment on the stones 
Simulium larvae are not often found m numbers. This may perhaps be due to 
the growth affording shelter for predaceous insect enemies. It is also very noticeable 
that Simulium larvae are never very abundant in those stony streams where there 
are large numbers of a certain small caddis-worm which lives in pebbly cases attached 
to the stones. 

Larval Moults and Growth Changes. —\ have been able to follow the entire larval 
development in only one species, S. latipes, but it will probably be safe to regard this 
as typical of all in respect to the number of moults and the principal developments 
which take place in the larval stage. In this species there are six moults, the change 
to the pupa taking place at the sixth. The first-stage larva is readily distinguishable 
from the others by the presence of the egg-burster, a small chitinous tooth near the 
posterior end of the clypeus, it being much more strongly developed* in S. latipes and 
S. aureum than in S. venustum (2); in the former it is plainly visible under a 3” 
objective as a dark spot in the middle of the back of the head; in the latter it is only 
discernible with a higher power. The first-stage larva is also characterised by one 
or two pairs of small hairs on each side of the egg-burster , which disappear after the 
first moult ; by the smaller number of elements in the mouth-brushes and in the 


_. * The actual tooth is no larger, but is darker in colour and surrounded by a darker 
and more strongly chitinised area of the general integument of the head. In this respect 
Simulium differs strikingly from the CULICIDAE, in which the egg-burster is surrounded by 
a membranous area. 


214 F. W. EDWARDS. 


anal sucker—in S, latipes about 15 mouth-brush hairs and about 4 teeth in each 
sucker-row ; and by the apparently two-jointed antennae, the second joint being the 
minute cone-shaped one. The later stages differ from one another chiefly in the 
size of the head, and in the progressively greater number of hairs in the mouth-brush 
and teeth in the sucker-rows. ‘The full-grown larva of S. latipes has about 40 hairs 
in the mouth-brush and 12-15 teeth in the sucker-rows. The anal gills in this species 
are quite simple in the first three larval stages, very slightly branched in the fourth, 
and distinctly so in the last. In this species, as in several others, the head is much 
darker in colour in the younger larvae, and the markings are less distinct. 


Diagnostic Characters of Larvae and Pupae.—tit is not the writer’s intention to 
describe the early stages in any detail, but merely to indicate the main points of 
difference between them. In the larvae these are to be found chiefly in the number 
of joints in the antennae, in the form of the anal gills, in the markings of the head, 
and to some extent in the colour of the body. The structure of the mentum and the 
number of hooks in the anal sucker also vary much with the different species, and 
some use has had to be made of these pomts. The mandibles and maxillae certamly 
exhibit good specific characters, but these have been ignored, as the more obvious 
distinctions are sufficient to separate all the British species. In the pupae the only 
characters of systematic value are the form of the cocoon and of the prothoracic 
respiratory filaments, but these are often so well-defined that they form a much 
easier means of identifying the sect than any characters of the adult ; they have 
the advantage of being entirely uninfluenced by sex, and are subject to very little 
variation. 


Seasonal Varvation.—The occurrence of definite seasonal variation in the Diptera 
is only known in a very few instances. The only two cases, so far as I am aware, 
in which it is regarded as proved, have been recorded by Hensel* for Dryomyza 
flaveola, and by Aldrich} for Cerodonta dorsalis. 1t has also been suggested by Verrall 
that Tvpula oleracea and T. paludosa may be summer and autumn forms of the same 
species, but this is not definitely established. 


In the case of Simulium the spring brood of S. eguinum is conspicuously larger 
than the subsequent broods, while there may also be a slight difference in S. ornatum ; 
but this latter is such a variable species that it is difficult to say for certain. In 
S. argyreatum, however, we have a well-marked case of seasonal variation, as both 
males and females of the spring brood are fairly readily distinguishable by colour 
and markings, as well as by their larger size, from the later broods ; the spring form 
is described below as var. sericatum, Mg. For absolute proof that these two forms 
are really the same species it would of course be necessary to breed one from the other, 
which I have not been able to do, but still 1 think the assumption may be justified 
for the following reasons :—(1) the male genitalia are peculiar and are identical in 
structure in the two forms, which are also alike in all other structural characters ; 
(2) in any locality where one form occurs the other may also be found ; (3) the spring 
form is never found after about the middle of May, and the summer form never before 
the beginning of that month. 


* Berl. Ent. Zeit., 1870, p. 133. 
t Ann. Ent. Soc. Amer., xi, 1918, p. 63. 


ON THE BRITISH SPECIES OF SIMULIUM—IL. 215 


Blood-sucking Habits.—In my previous contribution I stated that some of our 
commonest species were probably not bloodsuckers, but this opinion has since 
proved to be quite erroneous, as I have had abundant evidence that both S. ornatuwm 
and S. latipes will attack man ; and though there are still species (such as S, awreum) 
against which there is no evidence, it will probably be safe to assume that all the 
species may on occasion develop this habit. It would seem that their blood-sucking 
propensity depends much more on the weather than on the particular species ; 
a still, warm and sunny afternoon in April or May, and perhaps to a somewhat less 
extent later in the year, will always rouse them to activity. Some further details 
on this subject are given in the sequel. 

Range of Flight.—It has often been observed that Samulium may be found at 
considerable distances from their breeding places, and though this may in part be 
due to some stream having dried up and so been overlooked by the observer, yet it 
is certain that these insects can fly long distances. I have frequently taken 
S. ornatum and other species at a distance of a mile trom the nearest possible breeding 
place, while on one occasion I took S. venustum on Birds Hill, near Clothall, Herts., 
two miles from its home in the river Ivel. This power of flight is not by any means 
confined to females in search of food, as most of my records of specimens taken far 
from their breeding haunts are of small swarms of males. 


Parasites and Associates.—E. H. Strickland* has described in detail three parasites 
of Simulium larvae in North America : a worm of the genus Mermis, the sporozoans 
Glugea spp., and an undetermined Gregarine. I have found what appear to be these 
same parasites in various species in this country, though the parasitised specimens 
are never in very large numbers, Mermis in particular being rarely met with. As 
observed by Strickland, parasitised individuals are generally much above the 
average size. : 


It may also be of interest to note that small Chironomid larvae are frequently 
found within the cocoons, particularly of S. ornatum. I believe these are the larvae 
of a small species of Orthocladius, the cocoons of which are often to be seen lying 
close against the side of a Simulaum cocoon. Whether the Chironomid is dependent 
upon the Simulium or whether it also occurs away from them I have not been able 
to ascertain. 

Simulium as Food for Fish.—From the immense numbers in which larvae and 
pupae of Simulium occur in many rivers, it might be imagined that they would form 
an important part of the diet of fish, but in the case of trout at least this does not 
seem to beso. I have examined the stomach-contents of a number of trout obtained 
by Mr. M. E. Mosely, and found very few Simulwm larvae or pupae among them. 
In so far as the genus was represented at all, it was mostly by adults (chiefly males) 
of S. ornatum and S. equinum. Mr. Mosely suggests that this would be accounted 
for by the habits of the trout, which feeds mostly at the surface or by suction from 
the bottom. The adult flies found would be those which had fallen into the water ; 
among these, males from dancing swarms would no doubt be in excess. As noted 
below under S. tredecomatum, one case has been observed in which large numbers 
otf Semulwum larvae were found in the stomach of a trout. 


* Jl. Morph., Philadelphia, xxiv, 1913, pp. 43-105. 


216 F, W. EDWARDS. 


Control Measures.*—As has been pointed out by several previous writers, the best 
method for reducing the numbers of Simulium is the cutting and removal of weeds in 
rivers and streams; to be effective this should be done in March. This method 
would not affect those species which live chiefly on stones, such as S. reptans, 
S. tuberosum and S. latvpes. Probably all that could be done to check these species 
would be the removal of the most heavily infested stones ; the densest colonies could 
easily be discovered, as they are usually to be found mainly at certain favourable 
spots, 


TABULATION OF SPECIES. 


Adult Males. 


1. Front metatarsi more or less flattened in a vertical plane ; front tibiae, and 
usually also the thorax and abdomen, with silvery markings; claspers 
flattened dorso-ventrally (Group A) .. ai hes “s ae 

Front metatarsi cylindrical; front tibiae, thorax and abdomen without 
silvery markings, ee the tiviae may have silvery pubescence ; claspers 


not flattened (Group B) .. of uf as # ey i 
2. Hind metatarsi distinctly thickened in a vertical plane ; wna of thorax 
somewhat coarser ; claspers three times as long as broad ss Negi 
Hind metatarsi slender, nearly cylindrical ; pubescence of thorax very fine ;. 
claspers hardly longer than broad oe ae ae A si 

3. Hind tibiae on basal 4, metatarsi on basal 4, conspicuously pale .. ce ee 
Hind tibiae and metatarsi all black, or nearly so... at es nee 


4, Membranous area of pleura with soft hairs ; front metatarsi 5 times as long as 

broad ie ae : ie ar aig ..  ornatum, Mg. 

Membranous area of pleura Hie bare (even when freshly emerged); front 
metatarsi about 6 times as long as broad | 

5. Clasper rather slender, external margin slightly concave, the spine terminal 

varrvegatum, Meg. 

Clasper broader, external margin shila convex in the middle, the spine 


subterminal .. : ct monticola, Fried. 

6. Middle tibia» seo naaeiecialt scluut a at bes a be Ny yg aM: 
Middle tibiae all black .. ns ome tH cs. e a seer (BG 

7. Front metatarsi about 54 times as long as broad ; silvery markings of thorax 
inconspicuous ; stylus of adminiculum hairy... subornatum, sp. 0. 

Front metatarsi about 4 times as long as broad ; two conspicuous rounded 
silvery spots on thorax ; stylus of adminiculum bare .. reptans, L. 

8. Front tibiae conspicuously silvery in front .. ri oe ‘ye MP phi 


Silvering of front tibiae much less conspicuous ; clasper rather slender on 
apical half, with a dorsally-projecting thumb-like process near the base 
tuberosum, Ldst. 

* These are well summarised by Wilhelmi in a recent German Government publication 


(Die Kriebelmiickenplage ; Uebersicht iiber die Simuliidenkunde, besonders in practischen 
Hinsicht. Jena, Oct. 1920). 


10. 


11. 


12. 


13. 


14, 


ON THE BRITISH SPECIES OF SIMULIUM—II, 217 


Clasper shaped much as in tuberosum, but somewhat broader, and thumb-like 
process shorter ; adminiculum differmg from tuberosum and resembling 


the following ; } oe venustum, Say (=austem, Edw.). 
Clasper much broader, cine sub-basal thumb-like process and without 
articulated spine... j , oe .. morsitans, Edw. 

A pair of lateral, almost ilagigaligs nf oa heenils on front of mesonotum 
argyreatum (Mg.) Ldst. 


These markings greyish, not silvery and continued backwards as two narrow 
lines to the hind margin of the mesonotum 
argyreatum var. sericatum, Mg. 
Venation normal ; second joint of hind tarsi more or less excavated above near 
the base ; legs not densely hairy .. < is a iy iis rad, 
Venation abnormal, an extra vein (R,,,) present; second joint of hind 
tarsi cylindrical ; legs with long and rather dense hair 
hirtipes, Fries. 
Hind metatarsi almost cylindrical, and conspicuously pale towards the base ; 
membranous area of pleura with soft hairs ne equinum (L.) Edw. 
Hind metatarsi more or less thickened vertically ; eadios dark in colour ; 
membranous area of pleura quite bare .. ; ; ve ie . 
Pubescence of thorax golden ; second hind tarsal gine Sachi excavated. . 
Pubescence of thorax dull whitish aie second hind tarsal joint es 


slightly excavated ‘ e% subexcisum, Edw. 
Hind metatarsi barely three times as ae as ee vs latipes, Mg. 
Hind metatarsi quite four times as long as broad .. he we <a 
Clasper nearly cylindrical, of even width throughout .. angustitarsis, Ldst. 


reas much swollen towards the base, and rather short 
aureum, Fries. (= bracteatum, Coq. ; 
=angustipes, Edw.). 


Adult Females. 


Front metatarsi flattened in a vertical plane; front tibiae, and sometimes 
the thorax, with silvery markings ; last three abdominal tergites shining 
(Group. A}. . aby : : oa na 


Front metatarsi cylindrical ; front tihine and faves etait silvery markings, 
though the former may have silvery Riot ued last three abdominal 


tergites dull (Group B) vs td ri s. We. 
Frons (above antennae) and face (below paeeaadt orey, dull 23 HAMS, 
Frons blackish, shining (face usually grey) . zs “yf on 
Claws with a small sharp tooth just before the wiieitte xg = ge oe, 


Claws not toothed ; coloration like ornatwm ; membranous area of pleura bare 
subornatum, sp. Nn. 

Femora mainly dark; membranous area of Sad with soft hairs; front 
metatarsi 5 x1 a : ..  ornatum, Mg. 
Front and middle femora almost all Hatin! eee en area of pleura bare; 
front metatarsi 65x1 .. a . i varregatum, Mg. 


218 


10. 


nF 


12. 


F. W. EDWARDS. 


Claws toothed ; thoracic markings as in ornatum .. se a ae, | 
Claws simple ; thoracic markings different, or absent _.... me Boe fe 
Membranous area of pleura with soft hairs... ornatum var. nitidifrons, n. 
Membranous area of pleura bare e3 4 monticola, Fried. 
Face black, shining, like the frons ; legs were vided tuberosum, Ldst. 
Face grey ; legs more extensively pale by Me Ry .- NE 
Thorax almost dull, with coarser pubescence Le es a es 
Thorax somewhat shining, with much finer‘pubescence _... “if oe 
Basal two-thirds of hind tibiae clear yellow a i reptans, L. 
Basal half of hind tibiae Tne but less RY ao from the black 

apical portion ae ; iz wei ge 


Fig. 1. Head markings of Simulium larvae (the markings vary a good 
deal in intensity in most species; individuals have in each case been 
chosen which showed the markings most strongly developed. The figures 
are somewhat diagrammatic and not strictly to scale): a, S. ornatum ; 
b, S. variegatum and 8S. monticola; c, S. subornatum; d, 8. reptans ; 
e, S. reptans var. galeratum; f, S. morsitans ; g, 8. venustum ; h, S. argy- 
reatum ; 1, S. equinum; 39, S. latipes; k, S. aureum; l, 8. angustitarsis : 
m, S. subexcisum. 


Rather less than basal half of mid tibiae yellowish 
venustum, Say (=austen, Edw.). 
Rather more than half of mid tibiae yellowish sf .. morsitans, Edw. 
Thorax brightly shining ; front coxae reddish .. argyreatum (Mg.) Ldst. 
Thorax less shining, with traces of the two greyish longitudinal stripes seen 
in the male; front coxae blackish .. argyreatum var, sericatum; Mg. 
Vein R, ,, absent (normal); second joint of hind tarsi more or less excavated 
above near the base e ae : ae ee 3 


Vein R, ,. present; second joint of He tarsi eyindseal ; ; claws simple and 
rather small - east Tae ey ¥ me . hirtipes, Fries. 


13. 


14. 


15. 


16. 


17. 


ON THE BRITISH SPECIES OF SIMULIUM—II. 219 


Claws simple and very large ; thorax with three narrow black lines (hidden in 


good specimens by golden pubescence) A .. equinum (L.) Edw. 
Claws smaller and with a large basal thumb-like projection ; thorax unstriped 
14. 

Pubescence of sisi oiade or brassy ; second hind tarsal joint deeply 
excavated ; , Qe Ka ee 5 
Pubescence of thorax and a ea dull aiaeiahs -white ; second hind tarsal 
joint only slightly excavated .. “ d “r ag Fa 
Pubescence of abdomen very scanty, hairlike a aes abdomen pink in 
life; legs nearly all black .. +s ie .- latipes, Mg. 
Pubescence of abdomen dense, scale-like and ne abdomen yellowish in 
life “s «s ye oe oe - a aay |. 
Femora and tibiae clear yellow with black tips... .- aureum, Fries. 


Femora and tibiae darker, the latter with a dark ring close to the base 
angustitarsis, Ldst. 


Pubescence of thorax all pale .. ; .. .. subexcisum, Edw. 


Pubescence of thorax with three dark seectias “tie middle one divided by a line 
of pale hair-like scales... es :- es ap yerburyi, sp. n 


Full-grown Larvae. 


Antennae 4, 5, or 6 jointed, though the division between the first two, three or 


four is somewhat obscure a a or ay aa it 
Antennae 3 or 9 jointed a ; : ay “3 2 ee 
Last abdominal segment rounded Snail 6 or with only very small papillae ; 

antennae usually 5 jointed ae =o et és Lope 


Last abdominal segment with a pair of large maine papillae ; antennae 

4-jointed, but very long and slender .. a a us <u, Ae 
The three anal gills simple (sometimes with a few short branches in galerat:m) 4, 
Anal gills each with 5-10 branches* (4 or 5 in reptans) ae oe are 


~ 


Anal sucker with 100 or more rows of hooks, 18-25 hooks neachrow .. 5. 
Anal sucker with 65-80 rows of hooks, 12-18 hooks in each row ; body not 
bright green ; dark “ eyebrows ”’ present “5 ee ove s+ es 
Central and lateral teeth of mentum enlarged ; head markings distinct, but dark 
eyebrow absent; body bright yreen .. it -» equnum, L, 
All teeth of mentum smali ; head markings very indistinct, but a dark eyebrow 
present ; body colour uncertain .. Bh re tredecumatum, sp. n. 
Abdomen without distinct banding ; skin round anus with numerous minute 


trifid spines af } eas 


Abdomen with a distinct : Budee on dach peere dariall}: skin round 
anus nearly bare .. us Us ye a4 aH ar vs UB. 


* When the gills are ieianiticad thes are peer less frequently somemaneil at dian they 
aresimple. The gills of S. subornatum have not been observed ; since they were retracted 
in all the specimens examined, they are probably simple. 


220 F, W. EDWARDS. 


7. Head markings as in fig. la; anal spines very numerous and dark 
ornatum, Mg. 
Head markings as in fig. 1c; anal spines less numerous .. subornatum, sp. n. 
8. Back of head with a large roundish black area _.. reptans, var. galeratum, n. 
Back of head with four elongate dark spots, usually enclosed in a suffused 
brownish. area P a oe as argyreatum, Mg. 
9. Larger species; gills with 8-10 branches ; abdominal banding inconspicuous 
or absent ; dark “ eyebrow” usually conspicuous ; clypeal markings of 
the ornatum type ee often indistinct a also eee with 
different head marking) . Ge 


Lele abe 

6 [WW wrer">". 7 PAO™, 
fe N  aputeyn 
d Fl OG, h (tee 


Fig. 2. Apex of mentum of Simuliwm larvae ; 

all x 150. a, S. ornatum; b, S. subornatum; ec, 

S.reptans ; d, 8. equinum ; 'e, 8. latipes ; f, S. subex- 

cisum ; g, S. hirtipes (after Malloch) ; hs S. tredeci- 

matum. The hairs on the sides of the mentum are 

not shown, but are of diagnostic importance, as 
described in the text. 


\ 


ame ia, 
|| | 
l | b } d e f. 


Fig. 3. Antennae of Simuliuwm larvae; all 

x50. a, S. ornatum; b, S. subornatum ; 

c, S. equinum ; d, S. latipes; e, S. subexcisum; 

f, S. hirtipes (this last after Johannsen). 

Smaller species; gills with 4-6 (rarely 8) branches ; abdominal banding usually 
conspicuous ; dark “ eyebrow ” usually absent ; clypeal markings various 
but never of the ornatum type .. Ho std ne ey. i, 
10. Thoracic proleg with strong chitinisations behind the circle of hooks... 
variegatum, Mg. 
These chitinisations much less evident ; skin round anus with more numerous 
scales as oh fs ra by ies .- monticola, Fried. 


ki. 


12. 


13. 


14. 


15. 


16. 


ON THE BRITISH SPECIES OF SIMULIUM—II. 221 


Clypeus with a single dark spot posteriorly, or none iP x i=  .. 
Clypeus otherwise marked... wi Yr, we ie ar Pity OA 
A dark area in middle of posterior margin of clypeus, and another on each side 
of the head before the eyes ; body colour greenish. . reptans, L. 
Head all pale ; body colour greyish .. ds ig .. tuberosum, Ldst. 
A large black mark in the form of an H near the middle of the clypeus ; body 
colour yellowish-green .. SP on Pe .. morsitans, Kdw. 
A pair of dark spots on oduaned more or less connected anteriorly ; body colour 
bright green sit ..  venustum, Say. 
Gills branched ; clypeus with six a: ee not Aino well defined; ‘“‘ eye- 
brow ” without dark dot es ¥ em . latipes, Mg. 
Gills simple (at least in S. aureum): a dark dot in the anterior part of the 
** eyebrow ” re i. Ss a bee my a ad BS: 
Clypeus with a median dark streak, more or less interrupted, and four other 
spots aig sig Ee e™ be. os i aureum, Fries. 


Clypeus with eight distinct dark spots, including two transverse bars at each 
posterior corner ake 5a fe - angustitarsis, Ldst. 


Antennae very long, blackish, 9—joited ; central tooth of mentum simple 
subexcisum, Edw. 


Antennae short, pale at least ame 3- tes central tooth of mentum 
strongly trifid oe + a hirtipes, Fries. 


Pupae. 


td 
Prothoracic respiratory organs in the form of thick tubes, the main branches 
of which encircle the thorax at the nearly circular mouth of the 
cocdon 2 We - ve ne .. equinum (L.) Edw. 
Respiratory organs consisting of a number of slender filaments... sn. es 
Respiratory filaments with numerous ultimate branches (50-60 or more) : 
cocoon a loose shapeless network i af hirtipes, Fries. 
Respiratory filaments with not more than thirteen ultimate branches ; cocoons 
closely woven and shaped like the toe of a slipper, usually with semi- 


circular opening one ap = - os im Fl adits 9 
Respiratory filaments eight .. Wi i. - 2s Se i ae 
2 a SIX se at Te a i “he ts 9, 
_ os Tic 2. ~ es uh se - ees | 
s thirteen a as a tredecematum, sp. Nn. 


Cocoon with a long anterior dorsal projection : the upper two pairs of respiratory 
filaments with long vertical stalks .. - 2) yerburyt; spr Nn: 
Cocoon without anterior dorsal projection ; respiratory filaments otherwise 5. 


Filaments arranged in four shortly-stalked pairs .. via 8 
Filaments not in pairs ; all rather widely divergent slniee to the Sati towards 
which 3 of them are noticeably thickened . subornatum, sp. n. 


(713) C 


222 F, W. EDWARDS. 

6, Filaments much shorter than the pupa, all of equal thickness, the upper pair 
shorter than the others ; all four pairs with extremely short stalks : cocoon 
with ‘‘ windows” on each side in front 


reptans, L., and var. galeratum, n. 


Filaments about as long as the pupa; upper pair slightly thicker but no 
shorter than the others ; cocoon without ‘“* windows ” % ae 


Fig. 4. 
show branching; allx35: a, S. ornatum ; b, S. ornatum, var. ; ¢, S. ornatum var. 
nitidifrons ; d, S. subornatum ; e, 8. reptans; f, S. morsitans ; g, S. variegatum ; 
h, S. monticola ; i, S. venustum ; 9, S. tuberosum; k, 8S. argyreatum ; 1, 8. latipes; 


m, S.aureum; n, 8S. angustitarsis ; 0, 8S. subexcisum ; p, 8. yerburyt ; q, S. tredeci- 
matum. 


Bases of prothoracic respiratory organs of Simuliwm pupae, side view, to 


7. The four lower filaments arising from a short common stalk at the base ; then 


with longer separate stalks, of which that of the lower pair is the longer 
morsitans, Edw. 
The three upper pairs of filaments separate from the lowest pair at the base 8. 


8. The stalks of the two lower pairs of filaments usually longer than those of the 
two upper pairs 


All four pairs of filaments with short stalks of about equal length 


ornatum var. mitidifrons, n. 


ornatum, Mg. 


ON THE BRITISH SPECIES OF SIMULIUM—II, 223 


9. Cocoon with a long anterior projection ; upper pair of filaments with a long 
vertical stalk ; lower filaments twice dichotomously branched from base 
subexcisum, Edw. 


Cocoon without anterior dorsal projection ; filaments otherwise .. 10. 

10. Filaments all widely divergent at the base, the dapat one mere vertical 
for a short distance _ argyreatum, Mg. 
Filaments not widely divergent at bie, all more or less in the same vertical 
plane, arranged in three shortly stalked pairs yy A Ova: 

11. Filaments somewhat divergent, all about the same thickness ; cocoon all closely 
woven, as long as the pupa ye ..  venustum, Say. 
Filaments scarcely divergent, the lower pair cae thinner than the upper 

12. 

12. Cocoon short, usually not extending beyond the middle of the thorax ; filaments 
longer than the pupa, directed straight forwards .. tuberosum, Ldst. 

Cocoon longer than the pupa; filaments shorter... . 4 ites 

13. Cocoon all closely woven ; upper pair of filaments considerably thicker than 
the lower fe ; ne *. os monticola, Fried. 

Cocoon loosely woven at hd sides in dant, upper pair of filaments not quite 

so stout ¥: oo af iE 4 ¥ .. varregatum, Mg. 


14. Filaments in two distinctly stalked pairs ; cocoon with anterior projection 
. latipes, Mg. 


Filaments all arising from a common base, ornearlyso .. ey gee 15 
15. Filaments very stout at the base ; cocoon with anterior projection (short or 
long) o Ye +. angustitarsis, Ldst. 


Filaments slender at the ee cocoon without anterior projection 
aureum, Fries. 


1. §. ornatum, Mg. (figs. la, 2a, 3a, 4a & Db). 

Larva.—General colour variable, usually greenish grey or dingy greenish-yellow, 
without special markings on the body. Head light yellowish-brown ; a median 
wedge-shaped dark mark near posterior margin of clypeus ; a spot on each side of this 
and sometimes a trace of a fourth anterior to the wedge-shaped mark ; usually also 
pairs of marks on each side at the posterior margin ; a dark stripe above and in front 
ofthe eyes. Antennae 5-jointed ; first joint usually a little shorter than second, which 
is often very imperfectly separated from the first and sometimes obscurely divided 
into two in the middle ; fourth longer than third ; last joint very short and conical. 
Mentum* with three teeth much larger than the others ; between the central tooth 
and each of the other large teeth are three small ones ; sides of mentum with four 
to six long hairs. Last abdominal segment without ventral papillae. Anal sucker 
with about 70 rows of hooks each containing 12-15 hooks about 10ulong. Anal gills 
3, simple. Skin round anus beset with numerous minute scales dark in colour, and 
plainly visible under a magnification of 100, split apically into a number of spines 
(3-5). 


* Often incorrectly spoken of as the labium. 
(713) a2 


224 F. W. EDWARDS. 


Pupa.—Respiratory filaments about as long as the pupa, on each side 8 in number, 
arranged in 4 shortly stalked pairs, the stalks of the two ventral pairs usually the 
longest ; ventral pair slightly thinner than the others ; all the filaments more or less 
in one plane (the vertical). Cocoon tough, closely woven, about twice as long as its 
breadth in front ; height and breadth about equal ; anterior margin thickened, but 
without any forwardly-projecting piece in the middle ; no floor to the front half of 
the cocoon, which has its mouth widely open, and entirely covers the pupa. 

Habitat.—Principally a river species, but occurs also in small streams, some- 
times even those of a temporary nature. Where water-plants are present, the 
larvae and pupae attach themselves to the plants, and show a preference for those 
with long narrow leaves, such as Sparganium and water-grasses ; Ranunculus is less 
favoured, Potamogeton less still; usually the larvae are found chiefly on the lower 
sides, the pupae chiefly on the upper sides of the leaves. In the smaller streams, 
where water-plants may be few or absent, the larvae and pupae are to be found on 
or under stones or on any suitable objects. The species is rarely met with in 
streams less than 4 feet in width. 

Breeding Season.—This species appears to breed in suitable localities all the year 
round, almost regardless of the season. Pupae have been collected in every month of 
the year except January, from the middle of February until Christmas. On one 
occasion (19.11.1917) fully developed pupae were collected immediately after a period 
of three weeks continuous frost ; these produced adults of both sexes a few days 
later. I took a newly emerged male in the open on Ist March 1919. There appear 
to be three main broods, in April, July and September. 

Material collected.—Larvae and pupae have been collected as follows :—Bucks: | 
R. Misbourne, near Denham, 10.ix.15.; Burnham Beeches, iv. 1916, a few in stony 
stream with S. latvpes. Middlesex: Harefield, 21.1x.15, in stony and weedy 
stream ; Pinner, 22—24.vii.15. Surrey: Beddington, near Mitcham, 31.vu.19 ; 
larvae in millions in river Wandle, pupae also numerous, but no adults seen; a few 
pieces of leaf (totalling one yard in length) of Sparganium simplex carried approxi- 
mately 500 pupae and 4,000 larvae, apparently all of this species. Devon: Larvae, 
and pupae numerous in many small streams in the valleys of the Dart and Teign, 
up to a height of 1,000 ft. ; also in the rivers Otter and Sid, and in small runnels 
on the sea-cliffs near Sidmouth, vi. 1920. Hants: New Forest, 1.v.1920, a few in 
small streams and in Beaulieu River, in company with large numbers of var. 
nitidifrons ; River Test (M. E. Mosely). Herts.: River Chess, near Chorley Wood, 
30.vi1.15; River Lea, near Hatfield, v.1916 and 12.11.1917, on Ranunculus ; 
Pegsdon, 21.iv.16, on grass in small stream in “ The Meg,” larvae very large and 
very variable in colour; Hitchin, 5 i.15, under stones in stream 2-3 inches deep ; 
Radwell, 24.v.20, larvae of all sizes, very variable in colour, also some pupae, on 
stones in small rapid streams ; R. Ivel, near Baldock, vi.1917, on Ranunculus, Spar- 
ganium, etc. Beds.: Barton, 21.x.16, under stones. Cambs.: Bartlow, Hinxton, and 
Ashdon, 2.x.15, on grass and willow rootlets in small streams, none on stones in 
stream bed; Wittlesford, 29.xii.15 and 3.vii.16, larvae on stones in small stream ; 
Stapleford, 28.iv.16, on grass at edge of small stream ; Shepreth, 19.1.17 and 14.1v.17, 
on Ranunculus and Oenanthe in small stream; R. Granta, near Hauxton, 7.iv.17, 
on sedge at edge of river; Cambridge Botanical Gardens, 1.x.15, one pupa with 


ON THE BRITISH SPECIES OF SIMULIUM—II. 225 


numerous S. awreum on tree-roots in streamlet. Suffolk: River Lark, near Milden- 
hall, 25.iv.16, in large numbers on various water-plants, chiefly Ranunculus. Kent: 
Bushyruff, Dover, 21.ii1.1898 (B. L. Hesse). Wates: Anglesey, 19.ix.15, on stones 
in small stream across field (C. B. Williams). Channels Is.: Guernsey, Christmas 
1910, all stages together (K. G. Blair). Scotland: near Edinburgh, 27.1x.15, on grass 
at edges of small stream flowing through fields; current flowing at rate of 6 yards 
in 10 seconds (W. Evans). 


Variation.—In some localities the larvae of S. ornatum are fairly constant, in 
others they vary greatly in colour. Thus among 4,000 larvae from Mitcham, roughly 
examined, no variation was noticeable, while specimens from Pegsdon, Herts., 
and Harefield, Mx., varied in general colour from almost white to dark greenish, 
blackish or reddish; occasional specimens have dark brown markings as in S. aureum. 
The head markings are usually distinct, but may be obsolete ; very dark specimens 
sometimes have the head almost all black. The larvae found in small streams seem 
to exhibit much more variation than those found in rivers, the greatest variation being 
shown by parasitised larvae. This may possibly indicate that S. ornatum is 
normally a river species, the variation being induced by the change of conditions 
to those found in small streams. 


The pupae exhibit considerable variation in the length of the stalks of the pairs 
of respiratory filaments, but only in one specimen, out of hundreds examined, was 
any divergence seen in the number of filaments. This specimen had seven filaments 
on one side, eight on the other. 


The adults vary in size, and as is the case in some other species, the spring broods 
seem to be larger than the summer ones, thgugh this is not always very marked. 
The male varies to a remarkable extent in the silvery markings on the thorax, but 
only individually ; a close study of the variation might however possibly indicate 
the existence of definite local or seasonal forms. One such form has been recognised 
in the New Forest, and is described below. Most males have the thoracic markings 
similar to those of the female, and no more conspicuous ; some have conspicuous 
rounded silvery spots much as in the males of S. reptans, etc., while I have seen one 
or two beautifully marked examples with the whole of the margin of the mesonotum 
silvery, as well as the usual spots. All these are structurally typical; the only 
other character which I have noted to vary to any extent is the amount of black on 
the hind tibiae. The chief variation in the female is in the amount of white on the 
hind margins of the abdominal segments, which is sometimes very conspicuous, 
outlining three blackish areas on each segment. To this form Curtis’ name 
trefasciatum will apply. It is evident that S. ornatum is at present a plastic species 
some derivatives of which have attained stability, while others are probably now in 
process of development and fixation. It is therefore not surprising that the American 
representative (S. pzscicidiwm) should appear to be specifically distinct. 


Blood-sucking.—Since my previous notes on this subject were published, I have 
had abundant evidence that this species is a blood-sucker, but it would appear not 
to bite except on warm and rather still days, chiefly in the afternoon sun. Mr. C. L. 
Walton sent me some specimens from Crosswood, Cardiganshire, which he took in 
August 1916 attacking horses about the nose and eyes. I have also taken S. ornatum 


226 F. W. EDWARDS. 


(Letchworth, 5.iv 18) on horses’ belles, and (Letchworth, 19.x.17) in numbers on 
calves’ bellies, some containing blood. Besides this I have frequently taken females, 
sometimes apparently gorged, in cowhouses. Mr. W. Evans has noted the species 
biting his hand. I have also myself received bites from them on my nose, eyebrow 
and hand (25.iv.16, 19.x.17, 15.x1.17, 5-18 iv.19). The bites were not particularly 
painful and left no after-effects ; most of them were inflicted while I was in a stooping 
position (e.g., when gardening), and I frequently noticed that the flies were more in 
evidence then than while I was standing. Perhaps they mistake a man stooping 
for a horse or cow, which would seem to be their normal hosts. 

heferences.—The early stages of this species have been described in more or less 
detail by a number of writers. The best figure of the pupa is that of Castellani 
and Chalmers (Manual of Tropical Medicine, Ed. i. p. 812) ; the figure is not named 
specifically as that of S. ornatum, but is quite unmistakeable. A larva and pupa 
of S. ornatum have also been figured by Kollar (Sitzb. k. Akad. Wiss. Math.-Nat. 
KI. i, pp. 92-107, pls. iu, 1848, under the name S. serrceum, Mg ) ; Griinberg (Die 
Siisswasserfauna Deutschlands, 2a, p. 107, figs. 128-130, 1910); Verdat (Naturw. Anz. 
alle. Schweiz. Ges. v, p. 65, 1822—as S. serzcea) ; Liebe (Zeits. Naturw. 82, p. 345, 
1910—larva only); Meinert (K. Danske Vid. Selsk. Skr. (6) ii, p. 373, 1886). 
The most recent writer is Friederichs (Zeits. f. angew. Entom. vi, pp. 61-83, 1919), 
who figures the male hypopygium and the pural filaments of this species as S. reptans; 
with this use of the name veptans I cannotagree. It isnot clear what species he would 
indicate as S. ornatum ; possibly the one described below as S. subornatum. 


la. S. ornatum var. nitidifrons, nov. (fig. 4c). 

Closely resembles S. ornatum, Mg., except that the frons of the female is shining 
and not dull ; the mesonotum of the female appears distinctly more shining than in 
the typical form ; and there is little or no white on the hind margins of the abdominal 
segments. The male does not differ perceptibly from that of S ornatum. Larva 
with the head markings rather more distinct. Pupa with eight branches to the 
respiratory organs, all about equal in diameter, arranged in four pairs, all almost 
sessile or with very short stalks of about equal length. 

This form was abundant in the New Forest at the beginning of May 1920; larvae 
and pupae were collected on different water-plants in the Brockenhurst River, the 
Beaulieu River, and some small streams running into the latter. In the small streams 
it was the most abundant form Specimens of the normal form of S. ornatum 
were also reared from the same streams, but these were comparatively scarce. A 
few females were taken on the wing at Gidleigh Park, 8. Devon, 2.v1.1920, but no 
larvae or pupae were found. Both sexes were reared from pupae found in company 
with S. ornatum near Shelve, Salop, vu.1920, on grass in a small stream. 
Dr. G. Enderlein has since sent me specimens of both sexes (identified as S. reptans) 
from Berlin; he also sent specimens, probably from the same batch, to 
Friederichs, who has described them as S. reptans. | 

This form bears a great resemblance to the Scotch and Welsh S. monticola; the 
latter is regarded as a distinct species, since it differs markedly from S. ornatum 
in the larvae, pupae and male hypopygium, whereas the New Forest form apparently 
does not. 


ON THE BRITISH SPECIES OF SIMULIUM—II. 207 


2. §. subornatum, sp. n. (figs. lc, 2b, 3b, 4d, 7b). 


Adult.—Male: Thorax black, with fine golden pubescence and double silvery 
markings in front as in typical S. ornatwm. Membranous area of pleura bare. 
Silvery markings on the sides of the abdomen not very conspicuous, but extending 
on to the 8th segment as well as the 6th and 7th (which is not the case in S. ornatum). 
Genitalia resembling those of S ornatum, but the stylus of the adminiculum (viewed. 
from beneath) is much longer and narrower, more like S. reptans, but distinctly 
pubescent. Front tibiae silvery; front tarsi thickened, the metatarsus about 
51 times as long as broad. Middle tibiae yellow at the base Hind tibiae and 
metatarsi practically all black or dark brown, the latter moderately swollen. 


Female: Face and frons dull grey. Thorax marked as in S. ornatum, but rather 
less distinctly ; pubescence of mesonotum finer. Abdomen with the first five 
segments dull, blackish, with white hind margins, the last three moderately shining, 
but rather less so than usual. Legs coloured practically as in S. ornatum, but rather 
less brightly ; middle and hind tibiae sometimes with traces of dark rings near the 
base, especially on the under-side ; claws simple. Other characters as in the male. 


Larva.—Head markings similar to those of S. morsitans, but the H is not so dark 
and much less distinct. Abdomen without distinct banding. Antennae apparently 
6-jointed, but the first four joints not very distinctly separated ; the first about as 
long as the second and third together, the second, third and fourth about equal in 
length. Mentum with 9 poimted teeth in the terminal row, the middle one and the 
one at each end larger. Sides of mentum with 5 or 6 long hairs. About 80 rows of 
hooks in the anal sucker, 12-15 hooks in each row. Skin round anus with a number 
of minute spines or scales, which are smaller, less numerous and less conspicuous than 
those of S. ornatum. No ventral tubercles on last segment. Anal gills retracted 
in all specimens examined. 


Pupa.—Respiratory filaments 8 in number, the upper four rather widely divergent 
at the base, not arranged in pairs ; the uppermost one nearly vertical for some 
distance ; the upper three arise from a common base or a very short common stalk ; 
the next one is somewhat isolated ; the remaining four are in two pairs with very 
short stalks ; the three filaments on the inner side of the bunch are rather distinctly 
thickened towards the base. Cocoon differs from that of S. ornatum in being more 
loosely woven, with “ windows ”’ on each side in front. 


Material collected.—I first recognised this species from two females sent me from 
Bayswater, Oxford, 15.vi.16 (A. H. Hamm); the same collector subsequently 
obtained a third at the same place, 8.vi.18. About 10 ¢ and 59 were received for 
determination in 1917 from Bulwell Hall Park, Notts., 21.vi.16. and 18.vii.16 
(J.W. Carr). Some of these were presented to the British Museum, and I designate 
one of the males as type. I myself found a small swarm of the males hovering under 
a lime tree at Norton Bury Farm, Letchworth, 16.vi.17, and again at exactly the same 
spot on about the same date in 1918 ; in each case all the members of this swarm were 
this species, although S. ornatum was swarming under another tree close by ; I also 
obtained a single male by the river Severn at Shrewsbury, 24.vii.1920. A scrutiny 
of the series of S. ornatum in the British Museum revealed a single male of S. sub- 
ornatum from Llangammarch Wells, Brecknock (Lt.-Col. Yerbury). Finally 


228 F. W. EDWARDS. 


Mr. M. E. Mosely has presented to the British Museum a number of pupae and a few 
larvae ; masses of the former were in a jar in Mr. Mosely’s possession collected by 
the late F. M. Halford and labelled “ Crickmere, 16.ix.1894;” Mr. Mosely also 
collected a few pupae himself in the River Test, Hants. 


The species must be very similar to the North American S. piscicadium, but that 
is described as having yellow legs. 


3. §. variegatum, Mg. (figs. 1b, 49). 

Larva.—General colour brownish or dark grey, without special markings on the 
body. Head usually dark at the sides above and below the eyes ; clypeal markings 
often faint, when present usually ill-defined, but arranged much as in S. ornatum. 
Antennae 5-jointed, the first as long as the next two or three together ; second some- 
times indistinctly divided in the middle, twice as long as the third ; third shorter 
than the fourth. Mentum with 9 teeth in the terminal row, all shghtly trilobed, 
the small lateral lobes pale ; middle tooth very large ; about 8 long hairs on each side 
almost in a straight line. Thoracic proleg with a pair of rather strongly chitinised 
plates above the ring of teeth. Last abdominal segment without ventral papillae. 
Anal sucker with about 85 rows, each with 12-15 hooks. Anal gills each with 8-10 
rather long branches. Skin round anus with numerous minute dark scales, composed 
of short rows of spines (5-8 in each) ; the scales are much smaller and less numerous 
than those of S. ornatum. 


Pupa.—Respiratory filaments about as long as the pupa, six in number, in three 
very shortly stalked pairs ; all the filaments more or less in the same plane ; upper 
pair rather thicker than the others. Cocoon very large, fully a third longer than the 
pupa, in texture close-woven behind and above, but with an open network on the 
sides towards the front ; this network often, but not invariably, continued round 
the front of the cocoon and enclosing the ends of the respiratory tubes, the cocoon 
then approaching the “ boot ’’-shape of S. equinum. 


Habitat—Swift stony rivers and streams, both larvae and pupae on the upper 
surfaces of stones where the current 1s strong but smooth and the water not too deep. 


Material collected—Larvae and pupae were found in great abundance in early 
June 1920 in South Devon: in the River Dart through Holne Chase, the River 
Teign between Fingle Bridge and Christow, and the river Yeo near Ashburton, 
associated with S. tuberosum and S. reptans but in much larger numbers. A female 
specimen hatched from a pupa collected in 1912 at Ewbank Scar, Kirkby Stephen, 
Westmoreland (C. B. Williams), in company with larvae and pupae of S. monticola, 
Fried. 

Blood-sucking.—In August 1916 Mr. C. L. Walton sent me females from Crosswood, 
Cardiganshire, which were taken in company with S. ornatum “ attacking horses 
about nose and eyes and on belly.” 

Variation.—There seems to be great variation in the colour of the larval head, 
some being much darker than others, 

Reference.—Friederichs has described the larva, pupa and adult of this species 
as S. venefica, sp. n. 


ON THE BRITISH SPECIES OF SIMULIUM—II, 229 


4. §. monticoia, Fried. (figs. 1b, 4h, 5a). 

Adult.—Resembles S. ornatum in all respects except the following -—Membranous 
area of pleura bare ; front metatarsi about 6} times as long as broad ; male genital 
claspers broader, the external margin convex in the middle ; frons of female blackish, 
moderately shining ; abdomen of female entirely black. From 8S. variegatum the 
main if not the only differences are the shape of the male claspers, and the shining 
frons and dark femora of the female. This description agrees with that of 
S. arcticum, Mall., described from the female only from British Columbia, but without 
definite proof the two can hardly be considered the same. 

Larva.—General colour dirty greenish-grey, with obscure darker abdominal 
banding (on first 5 segments). Head very dark at the sides, except just round the 
eye-spots ; clypeus dark on its posterior margin, in the middle with a short dark 
longitudinal mark in contact with the dark margin, and slightly anterior to this 
a pair of inconspicuous dark spots. Antennae as in S. ornatum. Mentum with 
9 teeth in the terminal row, the central tooth and the one at each side rather larger 
than the others ; sides of mentum with 8-10,long hairs. No ventral papillae on 
last abdominal segment. Anal sucker with about 100 rows, each containing 15-18 
hooks. Skin round anus as in S. ornatum. Gills each with about 8 branches. 


Fig. 5. Male hypopygium, seen from beneath : 
a, S. monticola ; b, S. subornatum. 

Pupa.—Respiratory filaments barely as long as the pupa, six in number, arranged 
in three pairs with very short stalks ; all the filaments in a vertical plane ; upper 
pair slightly swollen at the base and thicker than the other two pairs. Cocoon as 
in S. ornatum. 

Habitat.—Moderately small swift streams, usually on rocks or stones ; both larvae 
and pupae usually on the upper or down-stream side. Sometimes present on stones 
covered with moss. 

Breeding Season.—The first brood hatches in May, and there is certainly another 
brood, since a gravid female was taken near water in the middle of June, and full- 
grown larvae have been found in August. 

Materval collected.—The species was fairly common in most of the small burns in 
the north half of the Isle of Arran, 22.v.—7vi.19 ; there were few larvae left by the end 
of this time. A specimen was also taken in a wooded valley at Ffrith, Flintshire, 
10. vi.19, and in the stream near by were numerous small larvae, possibly representing 
a second brood of this species. One or two specimens were reared from pupae 


230 F. W. EDWARDS. 


collected on stones in a rapid mountain stream at Kirkby Stephen, Westmoreland, 
22.vii.1912 (C. B. Williams). Inverleithen, near Peebles, 7.viii.1910, adult males on 
the wing, larvae of various sizes and some pupae on stones and some on grass 
(Dr. J. Rettie, per Dr. J. H. Ashworth). Various streams in the neighbourhood of 
Pontesbury and Church Stretton, Salop, vu.1920. 
Blood-sucking.—No definite evidence, but a few specimens (of both sexes) entered 
tent. ; 

References.—Friederichs, in the paper cited in the introduction (Oct. 1920), 
describes this species from the Harz Mountains. I have distributed it to various 
correspondents under the MS. name obreptans. 


5. §. reptans, L. (figs. 1d, 2c, 4e). 

Larva.—General colour greyish. Head with a single dark spot of varying size 
and shape in the middle of the posterior margin of the clypeus ; no dark mark above 
eye-spots. Abdomen with conspicuous dark greenish bands on all the segments 
dorsally, those on 6-8 nearly confluent. No ventral papillae on last segment. 
Gills each with 4 to 5 short branches. Skin round anus with numerous minute 
almost colourless simple scales. Teeth of mentum rather blunt, the nine in the 
terminal row all slightly trifid, the small lateral lobes pale. Antenna 5-jointed, 
gradually tapering to the end of the 3rd; first scarcely as long as the next two together, 
second nearly twice as long as the third. About 75 rows of hooks in the anal sucker, 
12-15 hooks in each row. 

Pupa.—Respiratory filaments much shorter than the pupa, 8 in number on each 
side, spreading but little from the vertical plane ; arranged in 4 very shortly stalked 
pairs ; uppermost pair rather shorter than the others ; all the stalks equal in length 
and all the filaments of equal diameter, slightly and evenly tapering towards the tips. 
Cocoon as in S. ornatum, except that on each side towards the front there is a rather 
large open space, usually crossed horizontally by one or two bands of silk. 

Halitat—Swift shady rivers in mountainous regions, but not at high altitudes. 
Where water-plants (such as Ceratophyllum) are present the larvae and pupae occur 
on them in some numbers, but are most typically found on the flat upper surfaces 
of large stones where the current is strongest, and particularly in those places where 
the flow is smooth and even and where there is no great depth of water. No larvae 
were found in the bare valleys above the tree-line, and very few in places where the 
rocks were much covered with moss. 

Breeding season.—The first brood of the year does not appear until the end of May 
or the beginning of June. I have had no opportunity of ascertaining whether there 
is a second brood, nor what is the method of over-wintering. 

Material collected.—The species was abundant in Machrie Water and in the burn 
flowing through Brodick Woods, Arran, v—vi.1919, in company with S. tuberosum. 
It did not occur in the smaller burns on the island. Also found in Devonshire, 
in small numbers in the Upper Teign and in the Yeo near Ashburton, and more 
numerous in the Otter at Tipton St. John and in the Sid at Sidmouth, in the former 
case in company with large numbers of S. variegatum, and in the latter with 
S. ornatum and other species, 1-18.vi.1920. A single male was taken on the wing 
near Church Stretton, Salop, 21.vii.1920. 


ON THE BRITISH SPECIES OF SIMULIUM—II. 231 


Variation —None noticed, except, as already mentioned, the size of the single 
dark spot on the head of the larva. 


Blood-sucking.—I have little to add to my previous notes except that up to the 
time I left Arran (5.vi.19) I was not attacked, and no females were even seen on the 
wing in Devonshire (up till 17th June). Probably the insect requires to be on the 
wing for some time before proceeding to its blood meal. Mr. P. H. Grimshaw writes 
“The specimens from Rannoch repeatedly flew at my face, in fact chiefly at my 
eyelids, and proved very irritating. They did not actually bite, but possibly would 
have done if I had given them the chance.” 


5a. §. reptans var. galeratum, nov. (fig. le). 

Adults.—Not positively identified, but probably identical with S. reptans. 

Larva.—General colour light greenish, with darker but not very conspicuous trans- 
verse bands on theabdomen. Head yellowish, clypeus witha black spot inthe middle 
of the posterior margin, connected anteriorly with a large roundish black mark which 
occupies almost the entire breadth of the clypeus ; its edges are not very sharply 
defined and it has a small anterior emargination. A dark rim above and behind 
the eye-spots, sometimes entirely surrounding them. Antennae apparently only 
4-jomted, the first joint longer than the remaining three together, second about 
equal to the third (the division between what are usually the first two joints seems to 
have disappeared). Mentum as in S. reptans. The pigment-spots within the sides 
of the sixth and seventh abdominal segments are darker and more conspicuous 
than usual. No ventral papillae on last abdominal segment. Gills either quite 
simple or with one or two (rarely three) short branches. Skin round anus with a few 
minute colourless scales. About 65-70 rows of hooks in the anal sucker, 10—15 
hooks in each row. 


Pupa.—Apparently indistinguishable from that of S. reptans. The respiratory 
filaments as dissected from the fully developed larva are as in S. reptans, and the 
single pupa collected that can with certainty be referred to this species (owing to 
its still retaining the larval head-capsule) showed no obvious difference either in the 
pupa itself or in the structure of the cocoon from S. reptans. This pupa was 
unfortunately not isolated, and the adult issuing was lost among a number of 
S. reptans. 


Material collected—Larvae were abundant on stones and water-plants (Spar- 
gamum, etc.) in the river Otter at Tipton St. John, 8. Devon, 14.vi.1920, in company 
with many S. ornatum and S. reptans, and a few S. equinum and S. aureum. 


Although this species was the most abundant in the larval state, even more so 
than S. ornatum, no adults were reared from pupae which could be correlated with 
these larvae, all the specimens obtained being either S. reptans (in the largest 
numbers), S. ornatum or S. equinum. This would seem to suggest that the form 
under consideration may be only a local form of S. reptans which has developed 
distinct characteristics in the larval state only ; but against this it must be stated that 
a small number of larvae were obtained at the same time and place which agreed in 
every respect with the Scotch form of S. reptans as described above. Moreover, 
in the river Teign, and even in the Sid, two or three miles distant, only normal 


252 F, W. EDWARDS. 


larvae of S. reptans could be found. Since, however, the only structural difference 
between this form and S. reptans is in the simple or less branched gills, it seems on 
the whole most probable that it is a variety of reptans, and I have so regarded it 
pending further investigation. The name has reference to the cap-like black patch 
on the head of the larva, a most striking feature. 


6. S. morsitans, Edw. (figs. Lf, 4/). 

Adult.—Both sexes have now been obtained by breeding. The female is very 
difficult to distinguish from that of venustum ; there seems to be no constant difference 
in the colour of the base of the abdomen, as I previously thought, but this species 
may perhaps be separated (in the female sex) by the greater amount of yellow on — 
the middle tibiae. The only clear distinction in the adult, however, appears to be in 
the shape of the male claspers. 

Larva.—General colour rather pale yellowish-green without distinct darker 
markings. Head yellowish, the clypeus with a conspicuous and rather sharply 
defined black mark in the form of an H. Antennae 4-jointed, the second joint a 
little longer than the first, third scarcely as long as second, fourth minute. Mentum 
with 9 teeth in the terminal row, the middle one long, the one at each end very 
broad, intermediate ones indistinctly trilobed ; four or five long hairs at the sides. 
Last abdominal segment without ventral papillae. Anal sucker with 65-70 rows 
of hooks, about 15-20in eachrow. Skinround anus bare. Gulls with 5 or 6 branches 
all approximately equal in size. 

Pupa.—Respiratory filaments 8, about as long as the pupa, in four pairs, the 
two lower pairs slightly thinner than the upper, and arising from a short common 
stalk ; each of the four pairs also with its own stalk, that of the lowest pair distinctly, 
often much, longer than the others ; all the filaments in the vertical plane, not at 
all swollen towards the base. Cocoon as in S. ornatum. 

Halitat—Weedy rivers of moderate current in company with 8S. ornatum, 
S. venustum and other species. Not found as yet in small streams. 

Breeding season.—There may be two broods in the year, as adults have been 
obtained in May and July, but the May brood is certainly the more numerous. 

Material collected—Two males were reared from pupae found in the river Granta 
at Hauxton, Camb., 5.vi.15, on Scirpus in company with large numbers of 
S. equnum and S. argyreatum. A fair number of larvae and pupae were found on 
grass and Ranunculus at the edge of the Lymington River, near Brockenhurst 
Bridge, New Forest, 2.v. 20. 


7. S. venustum, Say (austeni, Edw.) (figs. lg, 42). 

Adult female-—A number of females have been obtained by rearing ; they appear 
to differ from those of S. morsitans as shown in the table. I have failed to discover 
a better distinction. 

Larva.—General colour bright green, the abdomen with rather indistinct bands 
of darker green. Head light in colour, with a pair of somewhat oval dark spots at 
some distance from the neck, lying close together, connected at or near their anterior 
ends by a transverse bar ; usually also a dark area along the posterior margin of the 
clypeus. (The areas which in most species are dark are here pale, with dark areas 


ON THE BRITISH SPECIES OF SIMULIUM—II. 233 


more or less surrounding them.) Antennae and mentum as in S. morsitans, the 
mentum however with a broader black terminal portion. Anal sucker and gills as 
in S. morsitans. Skin round anus with some very minute scales, inconspicuous 
_ even under a magnification of 300. 

Pupa.—Respiratory filaments about as long as the pupa, six in number, arranged 
in three distinctly stalked pairs, the stalk of the uppermost pair nearly vertical, 
slightly longer and thinner than that of the middle pair. All the filaments in the 
vertical plane at the base, slightly and evenly tapering towards the tips, bases not 
at all swollen. Cocoon as in S. ornatum. 

Habitat.—Weedy rivers of moderate current. 

Breeding Season.—The first brood appears at the end of April; we have yet no 
clear evidence as to whether there isasecond. <A very few adults have been captured 
in August, and Miss F. Collins collected eggs in May 1911 at a spot where S. venustum 
was abundant, but these may have belonged to some other species. 

Material collected.—Larvae and pupae : New Forest, Hants., 1—-3.v.1920 ; numerous 
on grass and Ranunculus in Lymington River at Brockenhurst Bridge, also a few 
specimens in small streams running into Beaulieu River. Pupae: West Moors, 
Dorset, v.1911 (Miss F. Collins) ; R. Granta near Cambridge, 28.iv.16, on Ranunculus 
with S. equinum ; Stapleford, Cambs., 28.iv.16, one on grass at edge of stream, with 
S. ornatum. Adults, additional localities : Horley, Surrey, v.15 (@. A. H. Bedford) ; 
Oxford district, v.16 (A. H. Hamm); Letchworth district, v.17, v.18, v. 19, often 
at considerable distances from the nearest possible breeding places, some males as 
far as two miles; New Forest, 1—3.v.20, numerous in various localities, males 
swarming, females entering tent ; Shrewsbury, 21.vu.20, one female. 

References.—The pupa described and figured by Otto Fabricius (Schriften der 
Berl. Ges. Nat. Freunde, v, 1784, pp. 254-259, pl. iv) as that of S. sericea, Linn., 
may possibly be this species. Fabricius found the pupae in July on Potamogeton 
lucens in Norwegian rivers. 

A pupa figured (though not named) by Vogler (Mitt. Schweiz. Ent. Ges. vii, p. 279, 
1886) is almost certainly S. austent. Vogler found it in company with S. eqguinum 
(which he also figures) on Ceratophyllum and other water-plants in the river Rhine. 

S. austent is so extremely closely allied in all stages to the North American 
S. venustum, Say, as figured by Malloch and Pomeroy, that it cannot be ranked as 
more than a local race. I have compared the genitalia of British specimens with 
those of a specimen from Spartanburg, 8.C. (A. W. Pomeroy) and can find scarcely 
any difference. Both males and females are otherwise indistinguishable, and there 
is no structural difference in the larvae or pupae. Even the head markings of the 
larva are of the same type, though most of the British specimens examined had the 
dark markings developed to a somewhat less extent ; examples could probably be 
found however to match Pomeroy’s figure exactly. Though structurally identical, 
the two races seem to have developed biological differences. Pomeroy states that in 
America S. venustwm female is abundant from early spring to late autumn in the 
ears of horses, and that there are three or four broods in the year. In this country, 
as mentioned above, there seems to be normally only a single brood, while no examples 
have so far been found in horses’ ears. 


234 F. W. EDWARDS. 


8. S. tuberosum, Lundstvr. (fig. 47). 

Larva.—General colour dingy greyish or greenish grey ; head pale above, without 
dark markings ; abdomen with dark dorsal bands on the first five segments, the first 
two sometimes interrupted ; segments 6-8 all dark above. Antennae as in S. reptans. 
Mentum with 9 teeth in the terminal row, all simple, the middle one and the one at 
each end considerably larger than the others. Anal sucker with about 80 rows, 
each containing 12-15 hooks. Skin round anus bare. No ventral papillae on 
last segment. Gills each with from 6 to 8 branches. 


Pupa.—Respiratory filaments 6, in three pairs, each pair distinctly stalked ; 
filaments much longer than the pupa and all directed nearly straight forwards ; 
the two ventral pairs appear to be given off from the main stem which terminates \ 
in the dorsal pair. Cocoon constructed as in S. ornatum, but smaller, both actually — 
and relatively to the pupa, rarely extending beyond the middle of the thorax. 

Habitat.—With S. reptans. 

Breeding Season.—As in 8. reptans. 

Material collected——The species was very numerous in Machrie Water and 
Brodick Wood Burn, Arran, v—vi.1919, in the latter much more abundant than 
S. reptans, though in the former S. reptans predominated to some extent. Also 
found in Devonshire, in small numbers in the river Dart at Holne Chase, and the 
river Teign below Fingle Bridge. The occurrence of this species and of S. reptans 
so far south is of interest, and was somewhat unexpected, but their association 
with hilly country is confirmed. 

Variation.—None noticed, except to a slight extent in the size of the cocoon. 


Bloodsucking.—No further data available. 


9. §. argyreatum, (Mg.) Lundstr. (figs. 1h, 44). 

Larva.—General colour rather reddish grey. Back of head with four distinct 
rather elongate dark spots, the posterior one darker than the others, usually all 
enclosed in a dark cloud ; dark eyebrow present, but without any included darker 
dot. Abdomen with a conspicuous dark band on each segment. Antennae as in 
S. reptans. Mentum with 11 teeth in the terminal row, the middle one and the 
second from each end somewhat larger than the others. Anal sucker with about 
80 rows, each containing 14-18 hooks. Skin round anus with a very few simple 
scales. Very small ventral papillae on last segment. Gulls simple. 


Pupa.—Respiratory filaments 6, about as long as the pupa, not arranged definitely 
in pairs, but all arising almost from a common base and spreading out considerably 
at the base in all directions ; the uppermost one a little thicker than the other five, 
its basal part approaching the vertical. Cocoon as in S. ornatum. 


Habitat.—Weedy rivers with only a moderate current, usually associated with 
S. ornatum or S. equinum or with both these species. The larvae and pupae seem 
particularly fond of the leaves and stems of Scirpus lacustris, and of other plants 
with ribbon-like leaves, though they are by no means confined to these plants. 
The larvae predominate on the lower sides, the pupae on the upper sides of the 
leaves. 


ON THE BRITISH SPECIES OF SIMULIUM—TII. 235 


Breeding Season.—The larvae of the first brood become full-fed about the end of 
March, the flies appearing early in April, or possibly in March in some cases. 
Development must be rapid after this as second-brood specimens have been taken 
as early asthe 25th April. On the other hand, first brood specimens have been taken 
as late as 9th May (A. H. Hamm). The main second brood, however, appears at 
the end of June, and seems to be the most numerous in individuals. There is 
certainly also a third and perhaps a fourth brood. I have so far been unable to 
discover the species in any stage in the winter. The oviposition has not been 
observed. 

Material collected—Larvae and pupae have been collected as follows :—Cambs. : 
River Granta at Hauxton, 5.vii. 15, on Scirpus lacustris ; 7.1v.17, on sedge at river's 
edge; Shelford, ivii.l5. Suffolk: River Lark, near Mildenhall, 30.ix.15 and 
25.iv.16, on Ranunculus and Spargamum. Herts.: River Ivel near Radwell. 
Beds.: Cardington Mill, 28.vi.15, on leaves of Scurpus and Sparganium. Hants. : 
River Test (M. E. Mosely). Mr. H. Garnett has also sent me specimens taken near 
Evesham flying in clouds over the River Avon, in which thousands of Simulium 
larvae were living. 

Variation.—None has been detected in the larvae or pupae, but for the reasons 
stated in the introduction I have come to the conclusion that these are two well- 
marked seasonal forms of the adult, the first representing the spring brood, the 
second the subsequent broods. These can be distinguished as follows :— 

Spring form (var. sericatum)*: Length of body 3-3°5mm. Markings of male 
thorax not silvery, only dull greyish; the two patches towards the front of the 
mesonotum are produced backwards as two rather narrow lines as far as the scutellum 
and the thorax might therefore be described as greyish with three broad black 
stripes. Female thorax only moderately shining ; when viewed from in front it 
shows fairly evident traces of two longitudinal greyish stripes. Front coxae of 
female black, grey-dusted. 

Summer and autumn form (argyreatum) : Length of body 2-25 mm. Markings 
of male thorax silvery, especially when viewed from in front, consisting of a pair of 
more or less triangular patches, with their apices directed backwards. Female 
thorax rather brightly shining, without a trace of longitudinal stripes. Front 
coxae of female reddish, grey-dusted. 

Blood-sucking.—¥urther experience only confirms the view that this species is a 
habitual blood-sucker. At Mildenhall, 25.iv.16, it was in great numbers and very 
troublesome to a distance of quite half a mile from its breeding place, biting both 
head and hands. Though, to judge from the relative abundance of pupae, this 
species was much less numerous than S. ornatum, by far the greater proportion of 
the bites were inflicted by it, and I also noticed that S. argyreatum commenced its 
attacks much earlier than did S. ornatum. Other records were obtained in the 
Letchworth district, e.g., 18.1v.1919, bites on hand and ear. These were chiefly if 
not all of the spring form, but Mr. A. H. Hamm has provided me with several records 
of the summer form biting in the Oxford district (23.1x.16, 6.v1.17). 


* M. Séguy notes in regard to Meigen’s type of 8. sericatum, “ Détruit, le débris qui 
reste répond bien a votre description 9.” He also states that the types of S. argyreatum 
and some others *‘ répondent bien & vos tableaux.” 


236 F. W. EDWARDS. 


References.—It is possible that S. argenteostriata, Strobl, as described by Corti 
(Atti Soc. Ital. Sci. Nat. liii., p. 192, and liv, p. 223, 1914-16) is the spring form of 
this species, but I have not seen Italian specimens. The pupal filaments have 
recently been figured by Friederichs (Zeitschr. f. angew. Ent. vi, p. 61, 1919). 


10. S. equinum (L.) Edw. (figs. 12, 2d, 3c, 6a, b & c). ) 

Larva.—General colour rather bright green, body-markings darker green, but 
not conspicuous. Head with 6 distinct dark spots, 4 arranged as usual in the form 
of a cross, the other two being on the posterior margin of the clypeus. No dark 
stripe above the eyes. Antennae rather slender, 4-jointed ; the second joint nearly 
twice as long as the first or the third, which are about equal in length ; fourth minute. 
Mentum with 9 simple teeth in the terminal row, the central one and the one at 
each end much larger than the others. About 100 rows in the anal sucker, each with 
20-25 hooks. No ventral papillae on last segment. Skin round anus bare. Anal 
gills simple. 


b 


Fig. 6. Simulium equinum, L.: a, pupal skin of a hatched specimen 
projecting from mouth of cocoon, x10; b, respiratory organ of left side, 
seen from the right, xX 25; c¢, variety of pupal respiratory organ, X 25, from 
a specimen found in the River Test by Mr. M. E. Mosely. 
Pupa.—Respiratory organs consisting of thick tubes instead of long thin filaments, 
Close to the body of the pupa the organ divides into a dorsal and a ventral section, 
which are entirely in contact with the pupa on the one side and with the cocoon on 
the other. The tips of these dorsal and ventral sections are abruptly narrowed and 
shghtly overlap those of their fellows of the opposite side. From the dorsal section 
there project forwards five rather short and stout branches, two on the outer side 
and three on the inner ; the ventral section gives off a single branch near its base ; 
these branches are not more than one-third of the length of the pupa. Including the 
tips of the main sections, there are thus eight branches in all. The chitin forming 
the tubes is very thin and usually bears minute spines scattered over its whole 
surface, visible as dark dots under a magnification of 100. Cocoon closely woven, 
somewhat boot-shaped, that is to say, the anterior part of the floor is raised at an 
angle with the remainder, the opening being practically circular; anterior margin 
not thickened. 


ON THE BRITISH SPECIES OF SIMULIUM—II. 237 


Habitat.—Weedy rivers with only a moderate current, usually associated with 
S. argyreatum and sometimes also S. ornatum, on various water-plants, to a depth 
of 6-9 inches, the larvae mainly on the lower or down-stream side of the leaves, 
the pupae mainly on the upper or up-stream side. 


Breeding Season.—There are probably three main broods in the year, in March or 
April, July and September. No evidence has so far been obtainable as to the 
method of over-wintering, the earliest date on which larvae or pupae have been 
collected being 7th April, and the latest 10th September. Females, however, have 
been captured on the wing as early as February, and specimens of both sexes, 
obviously newly hatched, at the beginning of April. 

Oviposition.—Mr. H. Britten has given (Ent. Mo. Mag. May 1915) some interesting 
notes on the oviposition of this species, and m May 1919 I was able to confirm his 
observations, watching the females enter the water for the purpose of laying their 
eggs. In this case the eggs were deposited on a dead willow branch which was 
lodged in the river Ivel near Sandy, Beds, The insects chose a spot which was 
entirely sheltered from the current in order to enter the water, walked in to a depth 
of several inches and laid their eggs on the lower side of the branch. As Newstead 
noticed in the case of S. ornatum, the wings were wrapped round the abdomen and 
confined a layer of air between them and the body. Oviposition completed, the insect 
walked to the down-stream side of the branch, let go its hold and rose to the surface 
in its bubble of air. Some specimens took to flight the moment they reached the 
surface, others floated on the water for a few seconds before being able to rise. 


The eggs of S. equinum are not, like those of other species, enclosed in a hard 
gummy matrix, but in soft jelly like those of many CarronomipAE. They are 
however placed close together in a single layer. 

Material collected ——Larvae and pupae have been obtained as follows :—Bucks. : 
R. Misbourne, near Denham, 10.ix.15; R. Colne, near Denham, and stream in 
Denham village, 26.viii.15, larvae of all sizes and pupae, chiefly on water-plants, 
a few on stones. Herts.: R. Chess, near Chorley Wood, 30. viii. 15, immature 
larvae; River Lea at Hatfield, v.1916, on Ranunculus; River Ivel at Radwell, 
various dates. Cambs.: River Granta at Hauxton, 5.vui.15, in immense numbers 
on Scirpus and Potamogeton ; also at Shelford, 1.vu.15 and v. 17, and Grantchester, 
28.iv.16, on Ranunculus; Hauxton, 7.1v.17, numerous pupae and some larvae in 
one patch only on sedge at side of river. Suffolk: River Lark, near Mildenhall, 
25.iv.16, on Ranunculus, etc. ; numerous ¢¢ hovering in shade, but scarcely any 
2° seen on the wing, and none bit. Hants.: Bournemouth (D. J. H. Ashworth) ; 
River Test (M. EH. Mosely). Devonshire: In small numbers in the River Otter at 
Tipton St. John and in the River Sid at Sidmouth, mostly on water-plants, a few 
on stones. 

Variation —None noticed in larvae. In the pupae the minute spines on the 
surface of the respiratory organs are sometimes absent. The adults of the spring 
brood appear to be decidedly larger than those of the subsequent broods ; there is 
also much variation in the colour of the thoracic and abdominal pubescence of the 
females, some specimens being much more golden than others; possibly the spring 
brood might be distinguishable by being on the average less brightly coloured. The 

(713) D 


238 F. W. EDWARDS. 


dark stripes on the thorax of the female vary in width and distinctness, being 
hardly distinguishable in very dark specimens ; the legs are much darker in some 
specimens than in others. 

Examples fromthe Mediterranean region (e¢.g., those recorded in my previous paper 
from Fez) differ in the paler colour of the pubescence of the thorax in the female. 
Some pupae of this form collected by Capt. J. Waterston in Macedonia differ from the 
British forms in having the six minor branches of the respiratory organs longer, more 
slender, and closer together at the base. The larvae are alike. A single half- 
developed female pupa of this form has recently been given me by Mr. M. E. Mosely, 
who took it m company with normal pupae in the River Test, Hants., ix.1920 
(see fig. 6,c). 

Blood-sucking.—Ample confirmation has been obtained of the statement that this 
species normally feeds in horses’ ears, and it would seem that it is the only blood- 
sucking fly in Britain which does so; its activities must therefore be only too well 
known to agriculturists. Numbers of females were taken from ears of barge-horses 
on the canal near Uxbridge, 21.vii.16, no other species bemg seen. On 5th April 
1918 (as well as on other occasions) many were found in ears of farm horses at Letch- 
worth, from 30 to 40 in each ear, and though all were carefully examined no other 
species was found. Some of these specimens were so gorged that the blood was 
oozing from the end of the abdomen ; contrary to Pomeroy’s observation regarding 
S. venustum in America, they were not at all readily disturbed. On another occasion 
they were found to be much more numerous still, covering the inside of the ear like 
velvet, with their bodies closely packed, at right angles to the surface of the ear, 
and holding on only by their mouth-parts and front legs. Again at Digswell, Herts., 
9.vii.19, swarms of females of this species (and no other) were found flying round 
horses’ ears; the animals were much troubled by them and endeavoured to escape 
by standing in the river with their heads under the arch of a bridge, against which 
they could flick their ears. I have never found this species biting any other part of 
a horse. 


On one occasion (Letchworth, 19.x.17) a few specimens were found in calves’ ears, 
while at the same time there were numerous S. ornatum @& on the bellies of the 
animals. I think, however, there is no doubt that the horse is the main host, as I 
have several times searched for them in vain in the ears of cows. 

Mr. A. H. Hamm has supplied me with one or two additional records of the species 
biting the human subject ; I have not experienced the bite myself, though afew of 
the insects not infrequently fly round one. It is of some interest to note that, as 
might be anticipated, they fly round the top of the head and often settle on a person’s 
hat. 


References.—The remarkable pupa of S. eguinum has been described and figured by 
Vogler (Mitt. Schweiz. Ent. Ges. vii, p. 278, 1886), who did not however recognise 
the species he was describing. There are two other described pupae which bear 
an approximate resemblance to S. equinum. These are S. botulibranchium, Lutz, 
from South America and S. damnosum, Theo., from tropical Africa—the latter 
recently described by Pomeroy (Ann. Mag. Nat. Hist. (9) v, p. 80, 1920). The adult 
of Lutz’s species is unknown ; that of S. damnosum has a much greater resemblance 


ON THE BRITISH SPECIES OF SIMULIUM—II. 239 


to S. reptans than to S. equinum in general characters, though in the male genitalia 
there is certainly some resemblance between S. damnosum and S. equinum. From 
these and similar facts we may conclude that pupal characters may be useful as 
indicating unexpected relationships, but can hardly be adopted for defining 
subgenera. 


11. S. latipes, Mg. (figs. 17, 2e, 3d, 4/). 

Adult.—Rearing experiments have shown that the female I previously described 
as S. latupes was really that of S. angustitarsis; S. latupes Q is easily recognisable by the 
characters given in the key. The pink abdomen is nearly aprey very noticeable 
in life. 

Larva.—General colour dirty greyish or brownish, with indefinite darker markings, 
Head dark, with slightly darker markings arranged in the usual cross-shaped manner, 
but very indistinct. Antennae extremely slender, first joint about six times as long 
as broad, somewhat narrowed towards the tip, second nearly twice as long but not 
half as broad, third half as long and half as broad as the second, fourth minute. 
Mentum with 11 teeth in the terminal row, the central one and the second from each 
end much larger than the others. On the ventral side of the last abdominal segment 
are two conical papillae, nearly as long as the gills but with broad bases. Anal 
sucker of about 80 rows, each containing 12-15 hooks. Skin round anus bare. 
Anal gills each with about 8 branches. 

Pupa.—Respiratory filaments much longer than the pupa,four in number, arranged 
in two shortly but distinctly stalked pairs. All the filaments of about equal thickness, 
slightly and evenly tapering to the tips, and ‘arranged i in one plane (the vertical). 
‘Cocoon tough, closely-woven, about 1°5 times as long as broad, usually broader in the 
middle than in front, height in front barely equal to the breadth; front margin 
thickened and provided with a rather long median projection. There is a definite 
inner layer surrounding the abdomen of the pupa but not attached to the surface 
of the stone ; no floor to anterior part of cocoon ; mouth widely open. 

Habitat.—Small temporary streams and rills, especially those with stony bottoms, 
and perhaps most frequently in or near woods; only very exceptionally in rivers 
or fair-sized streams. The larvae congregate ee on the undersides of stones, or 
if these are not available, under dead leaves and twigs ; the pupae are more usually 
on the upper surface, and never deeper than an inch or two. 

Breeding Season.—The flies appear early in April, and there seems to be normally 
only one brood in the year; the rate of development of the larvae seems to be very 
irregular, and this will probably account for the presence of pupae as late as the 
middle of June. In favourable circumstances however it is not improbable that 
there may be a small second brood. 

Egeg-laying takes place in the early summer, and though I have never found the 
eggs, I have watched gravid females hovering over little sheltered spots of the rills 
and every now and then dropping suddenly almost to the surface of the water and 
rising again as suddenly. I could not ascertain whether eggs were being dropped, 
but if they were, the species has a very different method of oviposition from that of 
others of the genus. 

(713) : D2 


240 F. W. EDWARDS. 


Many of the streamlets in which S. latipes abounds are completely dry for the 
greater part of the summer, and though they may fill up during the autumn rains, 
I have not yet detected the presence of young larvae before about Christmas. It 
will be of considerable interest to discover exactly what happens to the insect 
during the dry season. I consider it most probable that it exists in the egg-stage, 
and that the egg-burster is a modification similar to that found in mosquitos of the 
Aédes group, which are capable of sustaining prolonged desiccation in the egg stage. 


Material collected——lLarvae and pupae occurred in abundance in most of the 
streamlets round Harrow, Pimner, Northwood, Ruislip and Stanmore, Middlesex, 
in the spring of 1915 and 1916. The earliest pupae were found about the end of 
March. Hatch End, Middlesex, 7.vui.15, larvae and pupae in stream which had 
been dry from middle of May to beginning of July ; larvae not nearly so common 
as in the spring. Streamlets in and near Knebworth Wood, Herts., half-grown 
larvae 4.1.17, larvae and pupae on stones and grass, 11.iv.17, numerous under 
stones, 10.v.20. Norton Common, Letchworth, Herts., small larvae xii. 1916, 
somewhat larger 1. 1917, pupae and some larvae 17.iv.17, a few larvae left 7.vi.17 ; 
many small larvae, chiefly second stage, but a few first and third stage, 21.xii.19 ; 
the stream was quite dry from June till early November 1919; larvae of all stages 
i-vi, but chiefly v, 29.u.20. Burnham Beeches, Bucks, iv.1916. River Lea, near 
Hatfield, Herts., v.1916, two pupae on Ranunculus in company with numerous 
S. ornatum and S. equnum ; one of these had the anterior projection of the cocoon 
forked, perhaps owing to its unusual environment. Streamlets on flanks of Goat 
Fell, Isle of Arran, v.1919, up to a height of 1,000 ft. Inverleithen, near Peebles, 
7.viil.1910, larvae of various sizes on grass (Dr. J. Retiie, per Dr. J. H. Ashworth). 
New Forest, Hants., 1.v.1920, larvae, mostly full-grown, and pupae, on grass and 
Ranunculus 1 small streams running into Beaulieu River, also in the Beaulieu 
River itself about a mile above Beaulieu. River Test, Hants, (MW. H. Mosely). In 
numerous small stony streamlets in the valleys of the Dart and Teign, S. Devon ; 
also a few in the River Teign below Fingle Bridge, vi.1920. 


Variation.—There seems to be but little variation in the adult or pupa, except to 
a slight extent in size and in the length of the stalks of the pairs of pupal filaments 
and the length of the anterior projection of the cocoon. The larvae also are fairly 
constant in any one locality, though the head markings vary in intensity, and the 
young larvae have usually darker heads than the larger ones, with less defined 
markings. The New Forest larvae had rather distinct reddish-brown bands on the 
abdomen, much as in S. aurewm ; these markings are usually hardly perceptible in 
Hertfordshire specimens. 


Blood-sucking.—For a long time I considered that this species was not a blood- 
sucker, and in fact it was long before I captured a female on the wing, in spite of 
searching for them near the places where the larvae were abundant. Swarms of 
males were found hovering in the shelter of hedges, but at first no females were seen. 
However in the Isle of Arran, at the end of May 1919, females were very numerous in 
Sannox Wood and by some of the wooded burns, flying round us in swarms, and so 
far as could be ascertained all the bites received were inflicted by this species; it 
could easily be recognised by the naked eye on account of the pink abdomen. 


ON THE BRITISH SPECIES OF SIMULIUM—II. 241 


References.—The larva and pupa have been figured (rather inaccurately) by 
Lt.-Col. A. Alcock in Entomology for Medical Officers, 1911, p. 126. There is also a 
very good and detailed description, with figures, of the tracheal system of both larva 
and pupa by Taylor (Trans. Ent. Soc. London, 1902, p. 701). Neither of these 
authors however had named the species. The species in which Weismann (Abh. 
Senckenberg. Natf. Ges, iv, p. 249, 1862) studied the development of the tracheal 
system would also appear from his figures to be S. latupes, not S. sericeum. 
Friederichs (Zeitschr. f. angew. Ent., vi, p. 61, 1919) sinks latipes, Mg., as a 
synonym of maculatum, Mg., but this seems to me to be without justification. 


12. S. angustitarsis, Lundstr.* (figs. 1, 47). 

Larva.—Head with eight dark spots above, four arranged in the usual cross- 
like formation, the other four in two transverse pairs near the posterior margin ; 
a dark stripe over the eyes, including a small blackish dot. Antennae much as in 
S. aureum, the second and third joints a little longer. Mentum as in S. aureum. 
The other characters cannot be given, as the only two larvae which have yet been 
found had partly transformed to pupae. The head-markings should be unmistake- 
able, though similar to those of S. aureum. 

Pupa.—Respiratory filaments four in number, longer than the pupa, all arising 
practically from the same point, widely divergent at the base, the upper two con- 
siderably swollen basally. Cocoon as in S. latipes, except that the anterior 
projection is usually very short. : 

Habitat.—Weedy rivers with only a moderage current, in company with other 
species, such as S. ornatum, S. argyreatum and S. equinum, but always nm much 
smaller numbers, and might usually be regarded as rather scarce. Very rarely in 
small streams. 

Breeding Season.—Nothing definite can be stated except that pupae have been 
collected from April to July and in September, while females have been taken on the 
wing in November. 

Material collected.—Pupae have been obtained as follows :—Cambs. : Stapleford, 
28.iv.16, on grass; l.vi.l5, on Spargamum. Herts.; River Lea near Hatfield, 
v.1916 ; River Ivel near Radwell, 16.vi.17, on Ranunculus. Suffolk: River Lark, 
near Mildenhall, 30.ix.15, and 25.iv.16, on Sparganium. Hants.: River Test 
(M. E. Mosely). Wales: Anglesey, 19.ix.15, one pupa on stone in small stream 
across field (C. B. Walliams). 

Additional localities for the adults are :—Timworth, Suffolk (Lt.-Col. Nurse) ; 
Wicken, Cambs. (F. W. E.); Shefford, Beds. (F. W. #.); Oxford district (A. H. 
Hamm); Woolhampton, Berks. (A. H. Hamm). Besides these, females were 
wrongly recorded as S. latvpes in my previous paper from Bovisand, Devon ; Corfe 
Castle, Dorset ; and Stockenchurch, Oxon. 

Variation.—Very little noticed, but the femora and tibiae of the female vary in 
ground-colour ; the lightest specimens show most distinctly the dark ring near the 
base of the hind tibiae, while in the darkest it is barely distinguishable. The hind 


metatarsi are always dark. 


* See synonymy below, and under 8S. awreum. 


242 F. W. EDWARDS. 


Blood-sucking.—No evidence. 

References and Synonymy.—The female I previously attributed to S. ee really 
belongs here, together with some of those I included with S. angustvpes, but the female 
I formerly regarded as this species is S. aureum. The pupa figured by Fries as 
that of S. reptans may be this species; also the pupal filaments figured by 
Témésvary (Rovartani Lapok, 1, p. 34, 1884) and Horvath (loc. cit. p. 195, pl. iii), 
though attributed to S. columbaczense, may belong to this species, as their structure 
seems identical. In any case the S. columbaczense of these writers would seem to 
belong to the same group as S. angustitarsis. Friederichs (Zeitschr. f. angew. Ent., 
vi, p. 61, 1919) found the species in the river Aar at Bern and figures the male 
hypopygium and the pupa and pupal filament; the figures have been transposed 
with those of S. maculatum (1.e., S. latepes). 


13. S. aureum, Fries* (figs. 14, 2m). 

Larva.—General colour dirty greyish or greenish grey, with rather ill-defined 
reddish-brown markings. Head with five or six well-defined elongate dark marks ; 
two transverse on the posterior margin, three or four longitudinal in the form of 
a cross, the middle one linear and sometimes divided (making six in all) ; all these 
sometimes enclosed in a dark cloud ; a dark stripe over the eyes, including a distinct 
blackish dot. Antennae with the first joint rather swollen, except on its apical 
fourth, which is rather suddenly narrowed, second joint much more slender but not 
greatly longer than the first, third half as long as the second, fourth minute. Mentum 
with 11 sharply pointed teeth in the terminal row, the central one and the second 
from each end much larger than the others. Two conical papillae on the ventral 
side of the last abdominal segment. About 70 rows in the anal sucker, each with 
12-15 hooks. Skin round anus bare. Gills simple. 

Pupa.—Respiratory filaments much longer than the pupa, four in number, all 
arising almost at the same point, but all practically in the same plane ; all equally 
though very slightly thickened towards the base. Cocoon constructed as in 
S. latipes, but without any trace of a median anterior projection. 

Habitat.—Small temporary streams, especially when stony; sometimes also 
found in rivers, but not in great numbers. 

Breeding Season.—My observations are somewhat contradictory and unsatisfactory, 
but there would appear to be at least two broods in the year, of which that appearing 
in the early autumn is much the most numerous in individuals. The only definite 
evidence available as to over-wintering is that Mr. A. H. Hamm took a female in 
the Oxford Museum, 3.11.1916. 

I found eggs, laid almost certainly by this species, in August 1915, in patches 
on blades of grass at the surface of the water, and on stones below the water. 
The eggs are dark brown and hard-shelled, not enclosed in jelly; some which I 
kept under a dripping tap hatched at the end of a week and the issuing larvae 
had well-developed egg-bursters, resembling those of S. latipes. 

Material collected.—Larvae and pupae have been collected as follows :—Middlesez : 
in all the small streams round Harrow, Pinner and Northwood, viii. 1915; most of 


* See synonymy below. 


ON THE BRITISH SPECIES OF SIMULIUM—II. 243 


these streams had contained S. latipes in the spring, but in August only S. aureum 
with (in one case) a few S. latipes and an occasional S. ornatwm could be found ; * 
in the spring of 1916 S. latipes was again abundant, and one or two S. awreum were 
found with it. Harefield, 19.ix.15, one pupa in small weedy stream with many 
S. ornatum. Bucks. : stony stream at Burnham Beeches, 10.ix.15. Devonshire : 
one pupa in a small stream at Start Point; one larva in the river Otter at Tipton 
St. John, vi.1920; one or two larvae and pupae in the river Sid at Sidmouth, 
vi.1920. Hants.: Botley, New Forest, i. 1915, among water-weeds (Lady Jenkyns) ; 
River Test (M. E. Mosely). Cambs.: Cambridge Botanic Gardens, 1.x.15 ; larvae 
and pupae in large numbers on roots of a tree in small stream 10 inches wide by 1 
inch deep ; all these had disappeared by Christmas 1915, and at Easter 1916 there | 
were no signs of larvae; very young larvae were present in June 1916. Suffolk: Barton 
Mills, 30.ix.15, a few pupae in a backwater of R. Lark; R. Lark near Mildenhall, 
25.iv.16, afew pupae with S. ornatum, etc., on water-plants. Lancashire: Manchester 
district (H. Garnett). Scotland: small rills on flanks of Goat Fell, Isle of Arran, v.1919 ; 
a few pupae among more numerous S. latipes. Additional records for the adults 
are :—Oxford district (A. H. Hamm); Porthcawl, Glamorgan (Lt.-Col. Yerbury). 
Blyth and Bulwell Hall, Notts. (J. W. Carr). Smailbeach, Salop (F. W. £.). 
Possibly a few of the records given previously for S. latipes really apply to this species, 
but both are common and widely distributed. 

Variation.—The central mark (or when it is divided, the two central marks) on 
the head of the larva varies somewhat in shape, the respiratory filaments of the pupa 
have sometimes a barely distinguishable stalk to each pair. The stilus of the male 
adminiculum varies slightly in width and the claspers in shape ; also the amount of 
hair on the basal pieces of the male genitalia is not constant. The male legs are 
usually almost entirely dark, but often the basal two-thirds of the femora and 
tibiae are perceptibly paler, this appearance being heightened by the golden 
pubescence, There isa variable amount of yellow on the hind metatarsus of the female. 

Blood-sucking.—No evidence. I have failed to induce captive females to bite by 
confining them over my hand, but this is also true of other species. 

References and Synonymy.—Rearing experiments have shown that the male and 
female which I previously described as S. awreum, Fries, really belonged to different 
species; I now propose to interpret Fries’ name as applying to the female described 
in Bull. Ent. Res. vi, p. 39, noting meanwhile that in that description I had overlooked 
the fact that the hind metatarsi are usually to a large extent yellow. The male 
I previously took for S. awreum therefore becomes S. angustitarsis, Lundstr. ; the 
male of S. aureum is really my S. angustipes (Bull. Ent. Res. vi, p. 40); this latter 
name therefore falls as a synonym of S. awreum, Fries. Most of the females assumed 
to be S. angustipes do not belong here but are really S. angustitarsis. 

S. aureum also occurs in North America under the name S. bracteatum, Coq. 
I have examined a male from Spartanburg, 8.C., 14.vii.13 (A. W. J. Pomeroy) and 
cannot find any appreciable difference either in the genitalia or in any other character. 

The larvae and pupae of S. bracteatum as described by Strickland (Jl. Morph. xxiv, 


form of S. latipes; but there are good structural differences between the two species in 
all their stages. 


244 F. W. EDWARDS. 


p. 45, pl. i, 1913) and Pomeroy (U.S. Dept. Agric. Bull. 329, p. 13) also agree almost 
entirely in structure, coloration and habits with British S. awreum, the only difference 
being in the shape of the larval antennae, which may depend on the preparation. 

The pupa has also been figured by Garnett (Trans. Manchester Microsc. Soc. 
1914 [1916], p. 10, pl. i) as that of S. reptans. The adults described by Corti (Atti 
Soc. Ital. Sci. Nat. lin, p. 192, 1914, and liv, p. 223, 1916) as S. awrewm apparently 
agree with the present interpretation of Fries’ species, which is evidently widely 
distributed in Europe. Dr. G. Enderlein has sent me specimens from the neigh- 
bourhood of Berlin; he also had determined themas S. awreum. There is an African 
representative with a wide distribution in that continent, which has recently been 
described by Pomeroy as S, aureosvmile. 


14. S. subexcisum, Edw. (figs. 1m, 2f, 3e, 40). 

Larva.—General colour pale dingy yellowish-brown. Head pale yellowish, 
clypeus with sharply defined black marks; the largest and most conspicuous a 
wedge-shaped one in the middle near the posterior margin ; a small one anterior to 
this ; a pair of small ones rather close together one on each side of the anterior end 
of the wedge-shaped one; also a transverse black mark at each posterior angle ; 
a conspicuous dark stripe over the eyes, including a small blackish dot. Antennae 
blackish, very long, 9-jointed, first jomt slightly swollen towards the base, about 
six times as long as broad, followed by four short joints, each a little longer than 
broad.; sixth and seventh rather longer, twice as long as broad ; eighth joint very 
long and slender, as long as the first; ninth minute. Mentum with 9 teeth in the 
terminal row, the middle one and the two at each end larger than the remaining four ; 
sides of mentum with one or two long hairs. A pair of well-developed ventral 
papillae on the last abdominal segment. Anal sucker with about 80 rows, each with 
about 10-12 hooks. Skin round anus bare. Gills simple. 

Pupa.—Respiratory filaments long, six in number; the first division is close 
to the base, into short stems diverging almost at two right angles ; the upper stem 
divides into two branches, which continue in a vertical direction as far as the height 
of the pupa, then bend forwards along the anterior projection of the cocoon ; the 
lower stem is shorter than the upper, dividing almost immediately into two branches, 
each of which is forked at some distance from its base. Cocoon as in S. latipes 
with a long anterior projection. 

Habitat.—Stones and grass-blades in small temporary streams in or near woods, 
in company with S. latvpes, but always comparatively scarce. 

Breeding Season.—The few specimens reared were obtained in the late spring. 
As in the case of S. latipes, there cannot normally be more than one brood in the year, 

Material collected—Two males were reared from pupae found on stones in a small 
rill on Stanmore Common, Middlesex, iv.1916. A single female was reared from a 
pupa found on a stone in a small stream at Knebworth Wood, Herts., 10.v.1920, 
and half a dozen larvae were found at the same time and place on stones and grass- 
blades. In each case the specimens were associated with large numbers of S. laivpes. 

A small number of females were taken at Glen Sannox and Glen Catacol, Arran, 
v.1919, flying in company with S. latipes ; also at Gidleigh Park, 8. Devon, 2. vi.1920. 


ON THE BRITISH SPECIES OF SIMULIUM—I. 245 


_ Variation —The pupal filaments dissected from one larva showed a small stump 
near the tip of one of the vertical branches, or both sides. The smaller dark spots 
on the larval head vary in size and distinctness. 


15. S. yerburyi, sp. n. (fig. 47). 

Adult female.—Differs from that of S. subexcisum, Edw., only in the colour of the 
vestiture of the mesonotum : instead of being all yellowish, there are three stripes of 
dark brown, the middle one divided by a line of pale scales or hairs; the middle 
stripe is longer than the side stripes, but does not nearly reach the front margin. 

Pupa.—Differs from that of S. subexcisum as follows :—Vertical stem of upper 
respiratory filaments shorter, forking near the base, each branch forking a second 
time at no great distance from the first fork, so that there are eight branches in all. 

Material collected.—A single female (the type of the species) was reared from a 
pupa collected at Knebworth Wood, Herts., 10.v.20, in company with many S. latipes 
and one S. subexcisum. There are three other females in the British Museum 
Collection, from Nethy Bridge, 18.vi.05, and Nairn, 28.v.05. (Lt.-Col. Yerbury). 
These were referred to in my previous paper as a variety of S. subexcisum, but since 
the difference in the female proves to be correlated with a difference in the pupa, 
it is perhaps preferable to regard the two forms provisionally as distinct species. 
The male and larva are unknown tome. ‘The discovery of the former may possibly 
prove the species to be identical with one of those described by Lundstrém. 


Sd 


16. S. hirtipes, Fries (figs. 2g, 3f, 7). 

I have not met with the early stages of this species, and they do not appear to 
have been described by any European author. Since however the American form 
described under this name appears to be identical in the adult, the following 
characters given by Johannsen and Malloch for American specimens will probably 
hold good for the British race also. 


Fig. 7. Simulium hir- 

tipes, Fries ; side view 

of respiratory organ of 

pupa (from Malloch). 
Larva.—Colour of upper surface yellowish on the thorax, fuscous on the abdomen. 
Head rich brown, posterior margin nearly black, Antennae 3-jointed,* first jomt 
occupying two-thirds of the length. Mentum with 7 teeth in the terminal row, 


* This may possibly have been an error of observation on the part of the American 
writers. The division between the first two joints is always rather indistinct. 


246 F. W. EDWARDS. 


all but the outer ones strongly trifid. About 100 rows of hooks in the anal sucker, 
12 inarow. Gills simple. 

Pupa.—Respiratory tufts much shorter than the pupa, divided near the base into 
four main branches, the two inner ones larger than the outer ones, each branch 
again dividing two or three times into twigs, so that upwards of 60 filaments may be 
counted. Cocoon, a dark matted mass of silk, of no definite form, secreted on the 
rock, the pupae only partly covered. 

Distribution.—Mr. F. Jenkinson has shown me a specimen said to have been taken 
at Crowborough, Sussex. Otherwise I have no records to add to those previously 
given, which all referred to the Scottish highlands. 


17. §. tredecimatum, sp. nov. (figs. 2h, 49). 

Adult.—Not yet recognised. 

Larva.—One of the largest, if not the largest, of the genus. Head markings of the 
same type as in S. ornatum, but more diffused ; a dark dot in the region of the eyes, 
and a pair about the middle of the clypeus. Antennae 4-jointed, the first two joints 
membranous, pale, the last two dark ; first and third joints about equal in length, 
second more than twice as long as either, fourth minute. Mentum with 11 divisions 
apically ; except for the middle three these are so short and blunt that they can 
hardly be called teeth ; the divisions are not noticeably trilobed. Sides of mentum 
with about four long hairs. About 110 rows of hooks in the anal sucker, and about 
18-23 hooks in each row. No ventral tubercles on last abdominal segment, but the 
whole segment is much more swollen ventrally than in most species. Skin round 
anus bare. Anal gills simple. 

One specimen was sufficiently developed to allow the pupal respiratory organs 
to be dissected out ; these agreed closely with the pupa described below. — 

Pupa.—Respiratory organ hardly more than a quarter as long as the pupa, 
divided into thirteen filaments in all. Atthe base there are four main divisions ; the 
first of these extends upwards nearly vertically for a short distance, emitting two 
branches close together near its base and a little higher up dividing into three ; 
the second divides into two close to its base; the third and fourth each split into 
three a little way from the base. No cocoon present in the material available. 

The above description has been drawn up from one pupa (evidently freshly formed). 
and a number of larvae in the British Museum, bearing the label, “‘ from stomach of 
a trout, England.”’ Most unfortunately no other data are available, but the species 
is so very distinct in both larval and pupal stages that it seems worth while to 
describe it, in order to call attention to the existence of such a remarkable form, even 
though it may ultimately be discovered that there was an error of labelling, or that 
the adult was already known under another name. ‘There is a bare possibility that 
the species may be S. hirtepes, the early stages of which, as mentioned above, have 
not been described from Europe. However it seems very unlikely that two forms 
which are to all appearances identical in the adult stage (as are the American and 
Kuropean races of S. hirtipes) should yet differ so widely as larvae and pupae. No 
species has hitherto been described from any part of the world with 13-filamented 
respiratory organs in the pupa. 


247 


CRAB-HOLES, TREES, AND OTHER MOSQUITO SOURCES IN LAGOS. 
By J. M. Dauzmt, M.D., B.Sc. (Public Health), F.L.S., 
Senior Sanitary Officer, Gold Coast. 


The occurrence of mosquitos in crab-holes is tolerably well known both in West 
Africa and in other countries. It is doubtful, however, whether these sources 
receive sufficient practical attention, and one hears too often when their presence is 
demonstrated the opinion expressed that the insects only shelter there and are not 
likely to use them as breeding-places when so many others are available. One has 
even found medical officers whose unbelief had to be assailed by demonstration of 
the larvae and pupae. Others again, though aware of the fact, perhaps think the 
occurrence of too trifling importance to justify the necessary steps to eradicate them, 
even when near a dwelling. 

The following notes from Lagos seem to support the opinion that where it can be 
shown that the imagines caught in houses are of the same species as those found as 
larvae in crab-holes in the vicinity, an effort to eliminate the latter is called for even 
at the expense of some labour and cost. In many cases the numbers issuing from 
a very few holes are so great that even if the species are not those known to be 
concerned in conveying disease, these efforts are Justified if it can be shown that 
the insects contribute appreciably to man’s discomfort, Reasonable comfort is 
one of the first conditions of health maintenance in the tropics, and the bites of 
mosquitos, even of those described—perhaps only in our present ignorance—as 
“harmless,” are amongst the many little but important matters which prevent 
both efficiency and enjoyment of life. 

The island of which the town of Lagos occupies the north-western portion is nearly 
flat, with a tendency to a whale-back contour in the occupied portion. The highest 
point is not much over 200 feet above lagoon level, and over the greater area water 
would be found at any depth from 3 to 10 feet, even at a distance from the lagoon 
margin. The soil is in the main sandy with areas of mud covered with coarse grass 
on many low-lying stretches towards the water-side. Being entirely surrounded by 
the comparatively still waters of lagoons and creeks its margins are at no point 
exposed to the surf of the open coast, and the rise and fall of the tide has only about 
a 2-footrange. The most familiar crab of the island is a land-crab, inhabiting in great 
abundance the low grounds near the margins of the lagoon at any point of the island, 
and travelling across country to any distance, restricted in the present instance by 
topographical limitations at most points to several hundred yards. This crab is 
Cardisoma armatum, Herklots, and is believed to be possibly identical with the 

West Indian C. guanhum. It belongs to the Family Grcoarcrnipas, land-crabs 
common to the warm regions of both hemispheres. The carapace is entire. It is 
very abundantly captured for food, being generally trapped on the pit-fall principle 
by means of a kerosene tin sunk flush with the surface in a likely spot, but being 
an inhabitant of filthy localities on Lagos Island its use for the table is not to be 
recommended. 


248 J. M. DALZIEL. 


(Another edible crab, Callinectes sp. (probably C. marginatus), is an article of 
food which is appreciated even by Europeans. It has a deeply serrated carapace 
with a longer spine at each end. ‘This is one of the swimming Crabs and is not 
concerned in the matter of mosquito breeding). 

Attention was first drawn to the connection between mosquito prevalence in 
certain houses and the propinquity of holes of the land-crab by observing the numbers 
of mosquitos which issued when the holes were disturbed. Collections of adult 
mosquitos made in the houses showed, in the second place, an almost exact cor- 
respondence with similar collections of adults issuing from holes in the garden or 
on neighbouring ground. Proof was then obtained by finding larvae and pupae in 
the water ‘of the crab-holes and hatching them out in the laboratory with ultimate 
determination of the species. 

Crab-holes may be found in almost any part of the island, but they are most 
abundant near the lagoon margins, where the salinity of the water tends to prevent 
their contributing to the mosquito pest. On the other hand, owing to the proximity 
of the subsoil-water to the surface all over the island, 1t may be that every crab-hole, 
except those daily washed by the tide, is at some time or other a potential source of 
mosquitos, provided that the water it contains is fresh or only brackish to a moderate 
degree and remains so for a sufficient number of days. 

On certain reclaimed areas in the town, which were originally creeks or inlets from 
the lagoon, they are particularly numerous. Such areas are generally overgrown with 
grass, and being at present without proper drainage they have acquired a spongy 
character, riddled with crab-holes and half-swampy in the rains. In these circum- 
stances Anopheline as well as other mosquitos breed freely, yet little except palliative 
measures can be taken until a more thorough reclamation with proper grading and 
drainage can be applied to alter the whole character of the locality. Their eradication 
is rendered the more important owing to the close proximity of native dwellings 
with a congested population, along with the observed fact that Stegyomyia fascoata 
does not shun the crab-hole, and the regular inspection of premises by the mosquito 
brigade doubtless tends to drive this domestic species to seek a less disturbed hatching 
place in the neighbourhood of houses. 

Although a considerable amount of routine labour has been devoted to these places 
in conjunction with general out-door work, most of the details recorded in these 
notes are the result of efforts directed against crab-holes in the vicinity of European 
Government Quarters near the lagoon and around the Golf-course at the eastern end 
of the inhabited town proper. In this area after attention was first drawn to their 
presence and potentialities it was found that a reasonable amount of properly directed 
effort was well repaid by results. 

A small gang of labourers with native overseers under supervision of a Huropean 
was detailed in May 1914 for out-door anti-mosquito work, and at first particular 
attention was paid to crab-holes in or near occupied European compounds. The 
part dealt with included Government House and various other residences of officials, 
along with the adjoining grassy slope of the Marina Road towards the edge of the 
lagoon, the Golf-course itself with the European compounds around it, and an 
unreclaimed marshy portion of the same area (here referred to as the Golf-course 
swamp). 


CRAB-HOLES, TREES, AND OTHER MOSQUITO SOURCES IN LAGOS. 249 


The method first employed was to pour down the hole a disinfectant such as 
cyllin, of strength about | in 8 or 10, with either cold or boiling water. The latter 
was found to be more effective in killing the crab and had the further advantage 
that cheap tar or crude creosote could be used. The boiling of the water on the 
spot, and the finding of dry fuel in rainy weather, however, involved so much. loss 
of time and waste of effort that a simple solution of cyllin or Kerol came to be 
preferred for use on a fairly extensive scale. (It may be said, however, that for 
private owners or occupiers troubled with a few crab-holes large enough to breed 
mosquitos, the occasional use of a bucket or two of boiling water mixed with 
Stockholm tar is a tolerably effective procedure. The crab either perishes in the 
hole or is dealt with in attempting to escape.) 

From 7th to 3lst May 1914, 7,250 crab-holes were treated, and 1,805 from 1st to 
15th June; total 9,055. After the middle of June on the other hand an accurate 
count was not attempted, as the rainfall brought the water near the surface, and in 
parts of the Golf-course area many holes lost their individuality in open pools and 
marshy spots. In fairly well-drained compounds they ceased to be found in more 
than small numbers where they had abounded a few weeks previously, and in many 
of these a periodic visit to deal with new holes was all that was required thereafter. 
As the number of compounds was considerable, and as crab-holes formed only one 
source of the mosquito trouble, it was found desirable in practice to make this work 
part of a regular routine, and as no means has so far been found to prevent a fresh 
access of crabs a certain amount of labour devoted to these continues to be necessary. 
The above figures therefore indicate the tentative work of the first few weeks. 

After the first efforts it was found that in a proportion of cases the crab survived 
and the hole reappeared in a day or two. Thus in a series of compounds 1,045 
crab-holes were dealt with by a first treatment with a disinfectant followed by 
obliteration of the hole. Of these 260 or 24°8 per cent. were found on a second visit 
to have reappeared. After a second treatment of these still a few persisted, and 
some of them had to be dealt with by other methods. In one compound situated 
within 100 yards of the lagoon edge, where various conditions were particularly 
favourable to the crab, over 1,000 holes were dealt with in the course of several visits. 

An objection to the use of tar or disinfectants in private compounds was found to 
be discoloration of grass or injury to plants, etc., and eventually the practice adopted 
for general use was simply to dig out the crab. Even by this method the creature 
occasionally defied discovery, either being absent, or, as proved by observation, 
having retreated to another gallery beyond the watery pocket at the bottom of the 
hole. 

The depth to which it is necessary to dig depends on the time of year and on local 
circumstances determining the distance of the water from the surface. The burrow 
is naturally oblique in direction, and a vertical depth of 3 feet has sometimes to be 
dealt with, the deepest measured being one or more in which larvae were recovered 
at nearly 4 feet from the surface. One such hole at the depth of 4 feet was followed 
for a further distance of 2 feet in a horizontal or slightly upward direction. It is 
obvious that mere obliteration of the hole without destruction of the crab is waste 
of time, except in virtue of removal for the time being of one mosquito breeding 
place, soon to be renewed. 


250 J. M. DALZIEL. 


A single hole may furnish eggs, larvae of all stages of growth, living pupae and 
empty pupal cases, indicating uninterrupted use by female mosquitos for several 
generations. The fact that in the great majority of cases the imagines captured were 
unfed and apparently quite recently hatched, supported the assumption that the 
insects were actually occupying the location of their birth. Other specimens are 
probably females returning to lay eggs. On one occasion several holes along the base 
of a line fence, which had been treated with tar emulsion a few days previously 
without obliteration of the burrows, were found to harbour adult mosquitos. On 
digging these out tarry water and tar-soaked soil were found but no larvae and no 
crabs either dead or alive. Probably the amount used had been insufficient and the 
creatures had succeeded in escaping through the soil from the end of the gallery or 
by another bolt-hole; the mosquitos were either dislodged individuals sheltering, 
or gravid females preparing to lay eggs. 

Conditions which interfered with the work of eradication were mainly due to 
the frequent occurrence of the holes along lines of fencing planted with shrubs and 
trees separating one compound from another, under clumps of banana trees, at the 
roots of ornamental plants, or at the edge of cement drains or masonry. 

Work on the above lines has been continued as circumstances demand up to the 
present time. Along the Marina in the main business street of the town numerous 
and very large crab-holes are found in which at times multitudes of mosquito larvae can 
be demonstrated. Probably the ultimate disposal of these will be through extension 
of the macadamised road surface to cover the whole space from the warehouses to the 
concrete sea-wall. 

In Bathurst crab-holes have been found to be a prolific source of Culicine mos- 
quitos. As digging in of the holes failed to stop the nuisance, the crab making a 
fresh outlet, the method adopted with success in some cases was to fill in with chloride 
of lime and stamp down (Annual Med. and Sanit. Report for Gambia, 1914, p. 13). 


Species of Mosquitos found in Crab-holes. 

In the years 1914 to 1918 inclusive numerous samples of larvae found in crab- 
holes were obtained and sent to the Medical Research Institute at Yaba.* There is 
no continuous record for all the consecutive months of any single year, and during 
the months of greatest rainfall, when more anti-mosquito work is done in the way of 
oiling and ditching on a wider scale, samples from individual crab-holes were Jess 
frequently sent, but the total number is sufficient to be of some value, and every 
month in the calendar is represented. 

On several occasions also adult mosquitos were captured in the holes for comparison, 
both with those found in neighbouring houses and with those ultimately hatched out 
in the laboratory from larvae collected at the same time. 

No note has been made of other inhabitants of crab-holes such as Copepoda and 
insect larvae other than CuLicipaE ; it is probable that species of Culicoides and 
other so-called “ sand-flies ”’ will be found amongst those enjoying the same habitat. 


* T have to thank the Director, Dr. A. Connal, for undertaking their investigation and 
Mrs. Connal for identifications of the numerous mosquitos obtained from these and from 
various other sources. 


CRAB-HOLES, TREES, AND OTHER MOSQUITO SOURCES IN LAGOS. 251 


(1) Mosquito Larvae in Crab-holes. 

From the samples sent to the laboratory, 14 species have been hatched. The 
following table gives the names of these and the number of occasions on which each 
was found. 


TABLE I. 

Species of Mosquito. No. of times found. | Percentage of total finds. 
Ochlerotatus irritans .. v6 Me | 95 34°6 
Oulex decens* .. . te bis 43 15:7 
Uranotaenia annulata as A 4} 15:0 
Ochlerotatus nigricephalus 22 8:0 
Stegomyia fasciata 20 7:3 
Anopheles costalis 2 bes 20 7°3 
Oulex insignis ce Me 13 4:7 
Culiciomyia nebulosa fa: 9 3°3 
Culex thalassius 4 1-4 

» Tima 3 Ly 

» salisburiensis .. 1 “4 

Uranotaenia bilineata var. fraseri ] “4 

Stegomyia luteocephala | 1 “4 

Micraédes inconspicuosus | 1 “4 
-| Total 274 


* Here and throughout this paper C. invidiosus is merged in C. decens. 


The number of samples is sufficiently large to ‘ive fairly representative results, and 
it may be accepted that O. zrritans is the species most often found. A further claim 
to priority in this respect is that it is not infrequently the only species found, and that 
it often hatches out in very large numbers from a single hole. Its occurrence in 
this habitat may be confidently expected in any month of the year, though perhaps 
the months of April, May and June have shown a preponderance of finds for this 
species. 

Its congener O. nigricephalus has the same seasonal range, but is far behind it in 
numbers. The former uses many other breeding places, but the only other common 
source for the latter has been surface pools, and once its larvae were found in a canoe. 
Both are vicious biters and are commonly caught in houses. 

Culex decens is well entitled to the second place. Its larvae seem to be less hardy 
than the others and are more difficult to hatch. The journey to the laboratory 
from the sources of collection is about 6 miles, and the agitation of the bottles used in 
conveying the larvae is perhaps more than some can stand. C. decens may be 
found at any time of year, but samples in the months of January to April have shown 
a greater proportion of this species. Its other habitats show almost as great a 
diversity as those of S. fasciata, and the adults have been found in dwelling houses 
in every month of the year. The third place on the list is taken by U. annulata, a 
species which has also been found in each month of the year, but which seems to 
have a special prevalence about January and February, when it often exceeds 
O. wrritans. Unlike the latter its adults are comparatively rare in houses except in 
the immediate vicinity of some prolific nidus. Its other sources are small swamp 
pools and wells. 


252, J. M. DALZIEL. 


Stegomyia fasciata and Anopheles costalis tie for the 5th place, but with two 
important distinctions, viz. (1) the occurrence of Stegomyia larvae has shown no 
regularity in number or in season of year, while Anopheles larvae have been found 
in crab-holes only in the period June to October ; (2) the localities of the respective 
sources (as recorded with each sample) have indicated for Stegomyra either the 
compound of a dwelling house (Huropean or native) or a place in comparatively 
close proximity ; in a few instances the Golf-course swamp is indicated, of which 
probably no part is much more than 200 to 300 yards from dwellings of some sort ; 
in the case of Anopheles the nature of the recorded notes, e.g., “‘in crab-holes full 
of rain-water,” “ along with tadpoles, etc.,” indicate along with the stated localities 
either a temporary meteorological condition or a permanent comparatively water- 
logged site as a constant accompanying factor. In many cases Anopheline larvae, 
often visible within the hole at a depth of a few inches, may have arrived there 
with the retreat of the water from the surface pool or swamp on which the eggs 
were laid. 


Culex insignis furnished about 5 per cent. of the recorded finds, but occurred in 
small actual numbers. No special seasonal incidence is suggested, and although it 
has been found in dwelling-houses, and perhaps oftener in out-houses, it is not a 
particularly domestic species. 


Culiciomyia nebulosa shares with S. fasciata the distinction of being the most 
domesticated of the mosquitos; indeed, its preponderance in organically polluted 
water or vegetable infusions in households probably gives it precedence in this respect 
in Lagos. Its occurrence in crab-holes is thus, as would be expected, less frequent, 
and generally in occupied compounds, though it has also been found in the Golf-course 
swamp. 


Of the remaining species in the above list one may say that C. thalassius as a lover 
of brackish water, though not dependent on it, is likely to be a more common 
inhabitant than here indicated of crab-holes in localities where this condition prevails, 
e.g., the tidal swamps at certain places on the island or neighbouring mainland at 
such a distance from the inhabited areas as to be beyond the radius of our special 
work. The species is common in houses in the vicinity of and to leeward of its 
breeding places, and bites freely. C. ima occurred in the vicinity of the Golf-course, 
and its adults were found in European houses, stables, etc., of the same locality, 
but no other breeding place has been observed. 


The 4 species at the end of the list are apparently exceptional. The larvae of 
S. luteocephala were associated in the crab-hole with those of S. fasciata and 
C. nebulosa, while about the same date the larvae of the first-named were found in the 
fork of a tree in the same European compound. Adults have been taken occasionally 
in houses of the neighbourhood. Those of C. salisburiensis have been found several 
times in surface pools, and the imagines at least once in a house of the same locality. 
No other source of the larvae of M. inconspicuosus and U. bilineata var. frasert 
has come to notice ; imagines of the former have not been captured, but those of 
the latter have at times shown a localised prevalence and at one time abounded in 
in the rooms of Government House itself. The occurrence of more than one species 
of larva in one crab-hole is common, and in fact perhaps almost usual, The two 


CRAB-HOLES, TREES, AND OTHER MOSQUITO SOURCES IN LAGOS. 253 


species of Ochlerotatus were often associated. ©. decens, C. nebulosa and 
U. annulata were all hatched from one sample. In another instance O. writans, 
O. nigricephalus, U. annulata and C. nebulosa were in association. 


(2) Adult Mosquitos in Crab-holes. 
A few collections were made of imagines found in crab-holes. The dates of these 
fall between the months of May and October, 1914, and representatives of 17 species 


were determined, viz. :— 


Ochlerotatus writans Culex fatigans 
- nogricephalus »  consemiis 
ri caliginosus Uranotaenia annulata 
punctothoracis ma biloneata var. fraseri 
Culex decens Culiciomyia nebulosa 
» insignis Mansonioides africanus 
,, thalassius Anopheles costalis 
» ruma Stegomyra fascrata 
salisburvensis 


The order of arrangement in the above list is not necessarily that of frequency 
of occurrence, and the number of individuals caught was insufficient for a true 
estimate of relative abundance of each species. A comparison of the two foregoing 
lists shows (1) that of the 14 species bred from larvae 2 have not been captured as 
adults in the crab-holes, viz., S. luteocephala and M. wnconspicuosus, of which the 
larvae were obtained only once each ; (2) that pf the 17 species caught in the winged 
state 5 are not represented in the list of larvae, viz., O. punctothoracis, O. caliginosus, 
C. fatigans, C. consimilis, and M. africanus. {these 5 could be regarded as occupying 
at the time of capture their place of hatching, the number of species definitely 
associated with crab-holes would be 19. The peculiar habit however of the larva of 
M. africanus in relation to the roots of the water lettuce (Pistea stratiotes) suggests 
its being only a temporary visitor after hatching in a pool or swamp near by (the 
locality was the Golf-course swamp and the month July in the rainy season). 

With regard to the other 4 species, the larvae of O. punctothoracis and of 
O. caliginosus have not been found in any of the collections from any source in 
Lagos during the period covered by these notes, but both have previously been 
obtained from borrow-pits on the mainland within the municipal area. 

The adults of both species have been caught occasionally in the houses of Europeans, 
and those of O. caliginosus were accompanied in crab-holes at a considerable distance 
from other possible breeding places by those of C. decens, which has already been 
shown to be one of the commonest species hatched from this type of nidus. 
C. consimilis has been bred from larvae taken from wells in Ebute Metta (on the 
mainland within the municipal area), and the adult appears not very infrequently 
in collections from the houses of European officials in Lagos itself. In crab-holes 
the mosquito was accompanied by adults of C. decens and S. fasciata, while all 3 
species were captured simultaneously in adjoining European quarters. C. fatigans 
has been hatched from larvae found in wells and barrels, and occasionally in a © 
boat or canoe. It is not a common mosquito in Lagos, but has been caught in 
European houses. It is probable that all 4 species occasionally make use of crab- 
holes as breeding-places. 

(713) E 


254 J. M. DALZIEL. 


Relation of Crab-hole Mosquitos to Human Beings. 

With reference to the possible influence of crab-holes on the prevalence of mosquito- 
borne diseases, the question will have to be decided on its merits in each different 
locality. In Tropical America two species, viz., Culex extricator and Deinocerites 

.sp., have been found to “ breed abundantly and exclusively in crab-holes,” but no 
evidence exists that either enters houses or shows any attraction towards human 
beings, nor was blood ever found in the females of the Culex examined (Knab, 
in ‘“‘ Science,” June 1910, p. 869). Crab-holes in mangrove swamps of Brazil were 
found to yield (along with CERATOPOGONINAE) Culex reticulatus, Lutz, C. corniger, 
Theo., and C. (Culicelsa) taennorhynchus, Wied. (Rev. App. Ent. i, p. 64), but there 
is no mention of their habits in relation to man. In the Panama Canal Zone crab- 
holes were dealt with in tens of thousands. 

According to the late Prof. Rubert Boyce, referring to Barbados, ‘“‘ In many of 
the more low-lying swampy coasts crab-holes occur in enormous numbers in the 
sandy soil, and in them are bred out vast numbers of mosquitoes. In fact they 
constitute the chief nuisance in those houses which are situated near the sea.” 
(“ Mosquito or Man,” p. 96). Howard, Dyar and Knab (“ The Mosquitoes of 
North and Central America and the West Indies,” p. 1) take exception to this and 
indicate that crab-hole mosquitos, as observed by them, do not molest man, and 
therefore their destruction need not be considered. It is apparent, however, that 
the species under consideration by these observers differ in habit from those observed 
by Professor Boyce, and this in turn is probably related to the important particular 
of special adaptation, and the authors’ conclusion must be read in the light of their 
statement that the mosquitos that breed in crab-holes are “ specialised forms of 
Deinocerites and related genera,” and that there also occur “a number of species of 
Culex peculiar to the crab-holes and even structurally these show more or less adapta- 
tion to their habitat.”’ (loc. cit., p. 149). 

In Lagos, of the species bred from larvae in crab-holes, all those which occur there 
in appreciable numbers have been found also in a variety of other sources. No 
species has yet been found here which shows a special adaptation structurally to this 
form of habitat. 

Three of the species mentioned in Table I have not, in connection with the present 
observations, been obtained as larvae from other sources, viz., C. rima, U. bilineata 
var. frasert and M. inconspicuosus ; of these the two last named were each found 
once only. It is unlikely therefore that the relationship in the case of these three 
has any importance from the point of view of endemic disease. But the crab-holes 

which produce abundance of possibly harmless species also furnish breeding-places 
for no inconsiderable numbers of those credited with the spread of disease. 

The probable effect of daily house-to-house inspection, with the main object of 
preventing breeding of domestic mosquitos, in driving the Stegomyra out of doors 
has already been referred to. In circumstances such as those in Lagos, therefore, 
the crab-hole in proximity to dwellings (a qualification which applies to practically 
the whole of the inhabited area) deserves attention as an accessory domestic breeding- 
place of the known vector of yellow fever. 

The connection of Anopheles costalis with crab-holes, I believe, on the other hand 
to be more or less accidental and dependent on the flat contour and generally 


CRAB-HOLES, TREES, AND OTHER MOSQUITO SOURCES IN LAGOS. 255 


low-lying and water-logged state of considerable areas of the island. It is doubtful if 
at any time of the year the female Anopheles finds it necessary to enter the shadow 
of a crab-burrow for purposes of egg-laying, at least to a distance beyond the range 
of ordinary daylight. War against the crab will therefore not be expected to have 
more than a subsidiary effect on the reduction of Anopheles, which must be dealt 
with by larger measures of drainage, oiling, etc., in view of the very numerous 
facilities afforded to that species on Lagos Island. 


Apart from S. fasciata and A. costalis, both carriers of Filaria bancrofti, the crab- 
holes do not appear in the present instance to be of particular importance in con- 
nection with filarial disease. 


In addition to supplying an extra and almost domestic source of S. fasciata (in 
itself a factor of sufficient importance) and to the possibly noxious—though still un- 
discovered—quality of some of the other species to which crab-holes afford unlimited 
breeding-places, thereremainsanadequate reason in Lagos forthe eradication of crab- 
holes in the mere fact of the numbers of mosquitos which they supply, of types 
which have been demonstrated as entering houses and preying upon human beings, 
even if they should be of secondary importance in relation to the spread of endemic 
diseases. It is quite a common occurrence for the sanitary gang to bring to light a 
sample of water from a single crab-hole containing many hundreds of larvae in all 
stages, and the ocular demonstration of the same to the occupier of adjoining premises 
has at times been adopted in the interests of propaganda, or to convince a complainer 
that the seemingly interfering methods of the workers are in reality well directed. 
From such a source when undetected amongst grass or under foliage, perhaps within 
a few feet of a dwelling-house, there may issue scores of mosquitos daily and there 
seems nothing to prevent this continuing for weeks or months as long as the burrow 
is occupied by the crab. There is unfortunately nothing to suggest that the land- 
crab includes mosquito larvae in its diet. 


A desirable consummation would be to keep the crab at a distance and prevent its 
access to the neighbourhood of dwellings ; but with a creature which exists in such 
numbers on the neighbouring sea-margin and which travels purposefully and with 
ease overland, up drains, etc., this seems at present impracticable, and some of the 
largest burrows may be found alongside a solid sea-wall or cement drain. 


Species of Mosquitos found in Tree-holes. 

Commencing in the rainy season of 1915 a regular search for breeding-places in 
trees was carried out. A ladder was used and sometimes a syringe with flexible 
rubber tubing. In a few months most of the trees in the public streets were dealt 
with, as well as many in private compounds. Hollows in tree-forks and holes in 
decayed wood or in cut branches were treated either by cement or other material, 
or by cutting out so as to prevent accumulation of water. In later seasons, so far 
as trees in public places are concerned, only an occasional routine inspection was 
required, but doubtless in private premises many escape attention. On only two 
occasions was a second find made in the same tree. 


In 1915, May to October, mosquito larvae were found on 43 occasions in 15 different 
species of tree or plant. For 1916 there is no record, but in later years others were 
(713) E2 


256 J. M. DALZIEL. 


found; e.g., in 1917, an exceptionally wet year, there were 20 finds in 14 species, 
and the total up to June 1919 includes 74 occasions in 27 different species. Rot- 
holes in cut timber are not included, nor are the rotting stumps of banana plants, 
which, when not rooted out, are a most prolific source of S. fasciata in native com- 
pounds. Samples of larvae were procured in a majority of instances, and the 
following six species of mosquitos were determined after hatching :— 


TaBie II. 
Species of Mosquito. No. of times found. Percentage. 
Stegomyia luteocephala. . ey 5) 33 61 
“ fasciata i th os 15 28 
Culiciomyia nebulosa .. ais sue 2 
Culex decens 30 4 a 2 | 11 
Ochlerotatus apicoannulatus .. i 1! 
Uranotaenia annulata . 1) 
54 


Apart from one or two exceptions from neighbouring villages within the municipal 
area, the Lagos samples were all from the town proper, including the native town 
and the area around the Race-course and Golf-course. The preponderance of 
S. luteocephala is obvious in this list and seems to be out of proportion when com- 
pared with tree-bred species in the Gold Coast and Sierra Leone. Other 
sources of this species in Lagos have been as follows :—Crab-hole (once only), a 
pool, the metal socket of a telegraph pole, and roof-gutters (similarly in Accra). 
In Freetown, Sierra Leone, its chief source was in hollows at the roots of the silk- 
cotton tree (Hriodendron orientale), occasionally in rock-holes, rarely in domestic 
utensils, and once in a coco-yam plant (Bacot—Report on Entomological 
Investigation undertaken for the Yellow Fever (West Africa) Commission, Aug. 
1914 to July 1915). 


The larvae of O. apicoannulatus have not been found in Lagos (during the course of 
these observations) except on the one occasion in a tree-hollow (amango). In Free- 
town, Bacot (loc. cit.) gives its sources as water-holes at the roots of Hriodendron 
and also rock-pools and occasionally tins. In Accra they have been obtained 
from a flamboyant tree in company with three species of Stegomyia (Ingram and 
Macfie, Bull. Ent. Res. vii, p. 145). A curious find in a banyan tree was 
S. luteocephala in a hole, and S. fasciata in a salmon tin lodged in the fork. In 
another banyan three species were found, viz., S. luteocephala in excess, along 
with S. fasciata and Culic. nebulosa. 

One species in the above list I do not remember having seen in records from 
others places, viz., U. annulata, found once alone and abundant in an ornamental 
screw-pine (Pandanus veitchii) at Government House. S. africana, along with 
Ochlerotatus marshalli, has been found by Graham in a cut bamboo, and 
O. longipalprs, Griinb. (S. pollinctor, Graham) in a hollow tree at Yaba (Bull. Ent. 
Res. 1, pp. 28-31). 


CRAB-HOLES, TREES, AND OTHER MOSQUITO SOURCES IN LAGOS. 257 


The commonest shade or ornamental trees planted or found growing naturally 
in Lagos are banyans (several species), mango, flamboyant, West Indian almond, 
Egyptian acacia and rain tree. Of the non-indigenous trees the flamboyant seems 
to be the most liable to rot-holes either in the fork or in the stump of a cut branch, 
and these holes generally retain water. The following list gives the names of the 
trees in which mosquito larvae were found; in two cases no record was made of 
the kind of tree by the finder. 


Taste ITI. 
Common Name. Botanical Name. Number of Occasions. 
Banyan (4 species) ft ..| Ficus thonningii 
»  platyphylla 22 
» populifolia 
» sp. 
Flamboyant aS oi ..| Poinciana regia | 13 
Mango “i ree 8 ..| Mangifera indica 6 
Pawpaw .. Carica papaya 5 
Woman’s Tongue or Egyptian 
Acacia .. Albizzia lebbek 4 
** Miraculous Berry (Yoruba 
—agbayun) sie ..| Sideroxylon dulecificwm 
Almond Tree ..| Terminalia catappa 
Black or Velvet Tamarind ..| Dialium guineense 
Rain Tree . aie oe ..| EHnterolobium duleis 
Breadfruit - eye ..| Artocarpus incjsa | 
Guava é ap a ..| Psidium guayava | 
Akee Apple ie ts ..| Blighia sapida 
Queen of Flowers mS ..| Lagerstroemia regina - 
(Yoruba—akoko) .. P ..| Newbouldia laevis 
Cashew... fe as ..| Anacardium occidentale 
ee a nS ..| Sterculia barteri 
Alexandrian Laurel “ ..| Calophyllum inophyllum 
Millettia sanagana 
Orange Citrus aurantium 
Cock-hat Tree (Yoruba—piregun) Dracaena fragrans 
Coconut . #. ..| Cocos nucifera 
Ornamental ‘Amaryllid wf Crinum giganteum 
Ornamental broad-leaved Screw- 
pine Pandanus pacificus 
Ornamental striped Screw-pine . ..| Pandanus veitchii 


The five species last named have not proved to be of as much importance as 
might be expected in Lagos with an average annual rainfall of about 75 inches, 
the number of larvae found under conditions likely to ensure full development 
being small. A notable absence from the list is the banana plant, which appears in 
some places to be a likely and important habitat. Possibly bananas have received 
less direct attention in Lagos than they deserve, but a good many trees have 
at different times come under my own observation without one positive find. 
On the other hand the stump left in the ground after removal of the decayed stem 
is an abundant and often neglected source of S. fasciata. The pawpaw is a source 
which is apt to be overlooked. The stem substance is soft and quickly decays, and 
a stump or cut branch containing a rotting hollow is soon concealed by the foliage 
of fresh young shoots. Also the stem of the healthy tree is hollow with incomplete 
partitions. A lateral perforation, caused probably by an insect, is frequently 
present and allows access to mosquitos. Sources of such a nature may remain 


258 J. M. DALZIEL. 


unnoticed for weeks or months and are commonly situated close to dwelling-houses. 
The vertical depth of the cavity in the interior of a pawpaw tree which was 
discovered to be the source of S. luteocephala and C. nebulosa captured in a 
neighbouring bungalow, was found on cutting it open to be 6 feet. 

The number of different species of trees harbouring either S. luteocephala 
or S. fasciata or both is considerable, and as regards the latter there is perhaps a 
predilection for banyans. Out of 33 occasions when S. luteocephala was identified, 
13 were from banyans (12 from the commonest species, viz., F. thonningi) and 8 
from the flamboyant. The occurrence of the different mosquitos in respect of trees 
was as follows: — 

S. luteocephala in banyans, 13; flamboyant,7; mango, 3; Egyptian acacia, 3 ; 
miraculous berry, 2; raintree, 1; velvet tamarind, 1; almond, 1; cashew, 1; pawpaw, 1. 

S. fascata in banyans, 6; flamboyant, 3; mango, 1; Egyptian acacia, 1; 
breadfruit, 1; Sterculia barteri, 1; coconut, 1; screw-pine, 1. 

C. nebulosa in banyan, 1; pawpaw, 1. 

C. decens in banyan, 1 ; flamboyant, 1. 

O. apicoannulatus in Egyptian acacia, 1. 

U. annulata in screw-pine, 1. 

Collateral observations on tree-breeding species in West Africa are referred to 
here for convenience of those interested in the subject. 

Sierra Leone (fide Bacot, loc. cit, suwpra).—In Freetown the “ Cock-hat”’ tree 
or “Cockade Bush ’* holds sufficient water in the axils and central leaf-whorl 
(in the same manner as the pineapple and other plants) to be a likely source 
of mosquitos. The breeding-place of two species in Freetown, viz. Uranotaenia 
ornata, Theo., and Eretmopodites dracaenae, Edw., seems to be restricted to this 
plant and to the similar conditions to be found in the coco-yam and the banana. 
Other species yielded by the “ Cock-hat”’ tree are S. simpsoni, and occasionally 
S. fasciata and C. nebulosa. 

The silk-cotton tree affords in hollows enclosed by the buttresses and amongst 
its spreading roots breeding-places for the following :—S. fasciata, S. simpsonc, 
S. luteocephala, O. minutus, O. apicoannulatus, C. nebulosa, A. costalis, and C. decens. 

The coco-yam (Colocasia antiquorum) yielded most commonly S. simpsoni and 
U. ornata ; also C. nebulosa and S. fasciata, and a few E. dracaenae. S. simpsont 
was found in banana plants, and S. fasciata in numbers with a few EF. dracaenae in. 
the sarsaparilla (svc). 

Gold Coast (fide Ingram and Macfie, Bull. Ent. Res. vii, 1916, p. 3; vin, 
1917, pp. 137, 145, 146).—The flamboyant has yielded most samples in Accra, but 
mango and other trees are also sources. Five species of Stegomyia, viz. S. luteoce- 
phala, S. fasciata. S. metallica, S. simpson, and S. unilineata, have been found, 
generally several species in association with each other or with O. apicoannulatus, 
Edw. Larvae of Cyathomyia fusca, Theo., were obtained from a flamboyant tree 

* This plant is an ornamental shrub of the Nat. Ord. Liliaceae, and is commonly 
planted along boundary fences in the coastal regions throughout West Africa. In 
Nigeria its Yoruba name is pireqgun and botanically it is Dracaena fragrans, though possibly 


other species may be included. Its popular names refer to the decorative use made 
of the leaves by the Sierra-Leonians on festive occasions. J. M.D. 


CRAB-HOLES, TREES, AND OTHER MOSQUITO SOURCES IN LAGOS. 259 


along with those of A. costalis, A. marshalli, S. fasciata, and S. luteocephala. 
In the Northern Territories of the Gold Coast, Ingram (Bull. Ent. Res. x, 1919, p. 50, 
et seq.) found amongst trees near dwellings that those which showed rot-holes most 
commonly were the flamboyant, wild fig trees, mangos, silk-cotton trees, “ dawa 
dawa ” (Parkia biglobosa) and tamarind. Larvae of S. fascoata, C. invidiosus, and 
S. ? unilineata were found ; also C. nebulosa at Kintampo in North Ashanti. 

In other countries, the larvae of Ochlerotatus (Culicada) ornatus, Mg., have been 
found in the horse chestnut (Aesculus hippocastanum) in Italy, and those of 
Stegomyva sp. in a tree fork and in cut bamboos at Hanoi, Tonkin. In the Lagos 
list no Anophelines appear, but in Freetown A. costalis, and in Accra both A. costalis 
and A. marshalli appear as facultative tree-breeding species. 


In India A. culiciformis, Cogill, and A. plumbeus, Stephens (syn. A. barianensis, 
James, and A. nigripes, Staeger) are believed to be essentially tree-hole mosquitos 
(Christophers, Ind. Journ. Med. Res. ii, pt. 3, Jan. 1916, and Christophers and 
Chand, zbid. pt. 4). The last-named, which has also been obtained from trees in 
Mesopotamia, freely enters houses and sucks blood. The related American 
species A. barberi, Coquillet, has a similar habitat. The larvae of A. plumbeus 
have been found in tree-holes both in England (Burnham Beeches, Liverpool, etc.), 
Scotland (Culross), and Ireland (Armagh, Derry, and Louth), in beech, horse- 
chestnut and elm. In Liverpool district and Cheshire A. plumbeus, either alone 
or in association with Ochlerotatus geniculatus, Ol., have been obtained in large 
numbers from tree-holes, the trees found most favourable to the insects being the 
elm, horse-chestnut and sycamore (Blacklock and Carter, Ann. Trop. Med. Parasit. 
xiv, pp. 115-126). In Edinburgh this species was found fairly abundantly some 
years ago in a well-wooded residential district on the south side of the town. The 
larvae were not found, but in the absence of any discovered source the suspected 
tree-holes were cemented up, with apparent reduction of the mosquito in the 
neighbourhood. 

Mosquitos breeding in Wells. 


In the period 1914 to 1918 inclusive, 12 species have been obtained as larvae 
from wells. The 265 samples from which determination of the imagines was 
made yielded results shown in the following table :— 


TaBLe IV. 

Species of Mosquito. Number of Occasions. Percentage. 
Stegomyia fasciata ene es 136 51°3 
Oulex decens oa at hi 39 14°7 
Anopheles costalis. . ie wt 35 13°2 
Ochlerotatus irritans 18 6°8 
Culiciomyia nebulosa 12 4°5 
Culex tigripes 6 2°3 

»  jatigans 6 oS 

» grahami 5 1°9 

» dutton 3 1] 

Uranotaenia annulata 2 7 

Culex consimilis 2 “7 

»  wnsignis 1 | “4 
Total 265 


260 J: M.. DALZIEL. 


The first five species in the list practically always occur in the same order in 
samples taken from wells, provided that a fair number are examined. C. decens 
favours nearly as great a variety of breeding places as S. fasciata. A. costalis 
tends to be relatively more frequent in wells located on vacant premises. C. tigrupes 
was always in association with one or more of other species, viz., S. fasciata, 
O. irritans, C. duttoni, C. grahami and C. fatigans, the larvae of the former preying 
on those of the latter. The larvae of C. consimilis have not been found (in the 
course of the work covered by these notes) except on the two occasions in wells, 
where they were also associated with S. fasciata. The adults have been captured 
in various houses of European officials and in crab-holes on the same premises. 


Mosquitos breeding in Boats and Canoes. 

A weekly inspection of boats and canoes around the foreshore is made and gives 
an opportunity for ascertaining the species which frequent them as breeding- 
places as shown in the list below :— 


Species of Mosquito. | Number of Occurrences. 


Anopheles costalis. . - _ a | 
Stegomyia fasciata a 
Culiciomyia nebulosa 
Culex thalassius 

»9 sitiens 

» fatigans 
Ochlerotatus irritans 

99 nigricephalus 

Anopheles funestus 
Culex duttoni 


ww 
a tet et ht DOD CO OI OD MI CO 


A. funestus was an unexpected find, and in spite of the existence of numerous 
apparently suitable conditions on the island it does not appear in any of my lists 
apart from this one, either as larva or adult. It is more abundant in fresh waters 
far from the sea. 


In a majority of cases the water in these sources is brackish, being lagoon water 
diluted by rain or vice versa. Under these conditions, although C. thalassius does 
not head the list, it is probably the most abundant survivor; e.g., in a canoe contain- 
ing distinctly brackish water, numerous larvae of C. thalassius and A. costalis 
occurred, but of the sample obtained, only one imago of the latter hatched out, while 
most of the former went on to full development. 


In the instances in which C. sitiens was found, the water was decidedly salt to 
the taste, and the imagines hatched out readily in the samples obtained. Analysis 
showed 1°8 per cent. estimated as chlorine, an equivalent of 2°99 per cent. sodium 
chloride. The larvae of this species have not been found in other sources during 
these observations. It has been recorded from Port Sudan in a pot sunk in salt 
water (Theobald, Mon. Cul. v, p. 388). 


CRAB-HOLES, TREES, AND OTHER MOSQUITO SOURCES IN LAGOS. 261 


The question of salinity in water samples harbouring mosquito larvae complicates 
the estimation of true prevalence of different species, some being more sensitive 
than others to variations of this nature. C. sitiens appears to be a true salt-water 
species, whereas C. thalassius is very adaptable, and though breeding abundantly 
in brackish marshes and lagoons such as those at Bathurst, Accra and Lagos, it is 
quite commonly found in perfectly fresh water. O. iritans seems to breed in 
brackish pools in the Accra lagoon with a salinity of 1°4 per cent. chlorine (=2°2 
per cent. NaCl). The following observation, however, suggests the need for dis- 
crimination as to the actual conditions under which larvae will not only remain 
alive, but proceed to full development. Very numerous larvae were obtained 
from two crab-holes close to the edge of the Lagos lagoon. The water was salt 
to the taste, with a salinity equivalent to 2°31 per cent. NaCl. Part of the sample 
was diluted with fresh water, while the rest was kept as obtained. From the 
former, O. wrritans hatched out abundantly for several days; from the latter 
numerous imagines of the same species hatched on the afternoon of the day on which 
the sample was collected, and on the following two days only, after which no further 
development of the numerous remaining larvae occurred. It is probable that the 
-crab-holes, originally containing comparatively fresh water, had been filled by 
a tide, and that the mosquitos which hatched on the first two days were those 
which had already pupated or which were ready to pupate at the time the sample 
was taken. In such a situation the water in a crab-hole may show many fluctua- 
tions in degree of salinity and tonicity, with their accompanying alternations of 
effect on the development of insect larvae, retarding the immature, but probably 
hastening the pupation of those far advanced.* 


‘Mosquitos breeding in Agbo Pots. 

The herbal infusions known as agbo are found in most native houses, especially 
where there are young children, and the receptacles are commonly concealed to 
escape the weekly inspection. They are very numerous, and sometimes the pots 
are of large size. On disturbing an agbo pot or calabash of long standing, the 
» number of insects which escape is almost incredible, and in many cases the fluid 
is a seething mass of Culicid and Psychodid larvae and pupae, along with the exuviae 
of their innumerable predecessors. From 450 samples of larvae from agbo the 
following species were obtained :— 


dl 

* Reference may be made to salt-water species occurring elsewhere :—Acartomyia 
(Aédes) zammittii, seems to live only in pools left after high water (Dardanelles). Stego- 
nocops sp. and Aédes fluviatilis (Culex fluviatilis, Lutz) ‘occur in rock-pools with concen- 
trated sea-water (Panama). Oulex sollicitans, Walk., is almost confined to salt pools 
‘near sea-beaches (Howard, Dyar and Knab, “‘ The Mosquitoes of N. and Central America 
and the W. Indies,”’i, p. 150). —C.. sollicitans, Walk., occurs in the mud of salt marshes 
(India, Karachi). Culex (Aédes) cantator, Coq., C. taeniorhynchus, Wd., C. salinarius, Coq.. 
with C. sollicitans, are found in the coastal salt marshes of America (Connecticut, etc.). 
Species of Taeniorhynchus and Anopheles occur in brackish marshes and mangrove 
‘swamps, and Anopheles subpictus (rossi) in brackish wells and salt-water tanks (India). 
A. (Pseudomyzomyia) ludlowi, a malaria carrier in the Andamans, breeds in salt swamps 
and mud flats. Anopheles multicolor, Camb. (Pyretophorus sp.) was found in Egypt in 
bho ae a 2°56 to 3:25 per cent. NaCl (Willcocks, Ann. Trop. Med. Paras. iii, 5, pp. 
586-587.) 


bo 
or) 
bo 


J. M. DALZIEL. 


TaBLE V. 
| 
Species of Mosquito. Number of Occasions. Percentage. 
Culiciomyia nebulosa .. sf “y 396 88 
Stegomyia fasciata.. oe a 51 1] 
Anopheles costalis si die ants 2) 
Culex decens me ue ‘ule i 1 
» ligripes .. Ji ie 4: 1 


Culex tigripes occurred in association with both C. nebulosa, and S. fasciata. 
S. fasciata occurs either alone or more often in association with C. nebulosa,. 
but the latter, apart from its preponderance in number of finds, undoubtedly 
furnishes immeasurably more imagines from this particular source than the former. 
This mosquito is exceedingly common in native houses and probably at present. 
greatly exceeds S. fascoata in numbers (see below). Although a considerable: 
number of ordinary water-pots are concealed or placed in locked rooms during 
the inspector’s visit, there can be no doubt that the weekly inspection of premises. 
is successful in detecting by far the greater number of domestic receptacles which. 
contain Stegomyva larvae, and thus disposing of large numbers of this species in 
its earlier stages. In addition to the agbo pot, apparently its most prolific source, 
C. nebulosa has been bred from samples obtained from the following :—catchpits- 
(for collecting household sullage), drains, water-pots, barrels, tins, a lawn-mower, 
roof-gutters, wells, pools, crab-holes, canoes, dung-emulsion (for smearing floors). 
tanning extract, starch emulsion, bottles oi palm-wine, and tree-holes. It will 
be noticed that several of these are of the nature of fluids rich in organic matter, 
in which, as in agbo particularly, the Psychodid, Pericoma meridionalis, Katon, is. 
its extremely abundant and almost constant associate. It may be added that 
C. nebulosa is by no means uncommon in pools away from dwellings, often in 
association with O. writans and A. costalis, so that domesticity, though a prominent. 
feature of its bionomics in Lagos, is not an essential characteristic. 


Mosquitos breeding in Roof-gutters. 


Previous to their gradual removal after installation of a pipe-borne water-supply, 
roof-gutters were fairly regularly inspected and swept. Comparatively few samples. 
have been examined, with the following record :— 


TasLe VI. 
Species of Mosquito. Number of Occasions. Percentage. 
Stegomyia fasciata : 1 56°5 
- luteocephala 17-4 


Ochlerotatus irritans 
Ouliciomyia nebulosa 
Anopheles costalis 

” mauritianus 


nt eet DO DO HE CO 


——— 
~) 
or) 
— 


CRAB-HOLES, TREES, AND OTHER MOSQUITO SOURCES IN LAGOS. 263 


The occurrence of the larvae of A. mauritianus, Grandpré, is unique in the present 
record. It is a very rare mosquito in Lagos, but the adults have been found at least 
once in the same locality, viz., the dwellings around the Golf-course. The larvae 
have previously been obtained at Yaba (which is within the municipal area) in a 
pond near the Research Institute, and in road puddles in the wet season. 


Mosquitos from various Domestic Sources. 

The weekly visit of the Sanitary Inspector has made it possible to obtain samples 
of mosquito larvae from domestic sources, and these have been collected whenever 
circumstances permitted of their examination at the Research Institute with a view 
to determination of the species. The following record is the result of material 
thus brought under observation during an aggregate of 45 months, between the 
years 1914 and 1918, thus :—1914, May to November, 7 months ; 1915, 12 months ; 
1916, January to April, and November to December, 6 months ; 1917, 12 months ; 
1918 January to August, 8 months. Amongst domestic receptacles are included 
for the present purpose, besides the usual utensils such as water or agbo pots, tins, 
bottles, tanks, barrels, etc., the following :—wells on the premises, roof-gutters, 
house-drains and catch-pits, but not puddles in the yard or crab-holes. 

No less than 17 species are represented, while the total number of species captured 
in Lagos during the period of the present record is 46. It may also be accepted 
that a collection under similar circumstances in any large coast town in West Africa 
will furnish a like preponderance of S. fasciata in greater or less degree ; ¢.g.,in 
Bathurst (date ?) with a collection obtained from 150 compounds each month, 
the proportions were Stegomyia 94 per cent., Culex 5°5 per cent., Anopheles 05 per 
cent. 

During two years in Accra, 1,121 domestic samples examined gave the following 
result (Ingram, Bull. Ent. Res. x, 1919, pp. 50-51) :— 

S. fasciata occurred in 876 or 78:0 per cent. 

C. fatigans es Nie ce 

A. costalis 4s oleae es ape 

C. decens 9 st Oe TOE AG 
C. wnvidiosus ,, SNe B aa eg he 

C. duttona i‘ eM: 5, ea eee 

C. tugrupes var. fuscus Bh) eB nips 

C. nebulosa ‘ tee ce te tos 

A collection in Lagos similar to the present one, composed of 1,043 samples and 
covering 8 months in 1910-1911, was recorded by Graham (Bull. Ent. Res. ii, 
p. 127) as below :— 


S. fasciata in 92°5 per cent. of domestic water-samples examined. 
Pectinopalpus fuscus* ,, 21°6 “5 ii ¥5 99 s3 
C. duttona sar OFS ‘3 a ‘b 0 2 
C. tagrvpes var. fuscus,, 5°3 ”: ¥ 2 . 9 
C. nigrocostalis * coo 28 \ ui ‘5 99 
Pyretophorus costalis ,, 18 ,, F a 99 29 


* Pectinopalpus fuscus, Theo. =Culiciomyia nebulosa, Theo. ; and Culex nigrocostalis 
Theo.=Culex decens, Theo. 


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CRAB-HOLES, TREES, AND OTHER MOSQUITO SOURCES IN LAGOS. 265 


In Lagos C. nebulosa is, next to S. fasciata, the species most common in domestic 
receptacles (greatly exceeding the latter in agbo pots), but in Accra C. fatigans 
apparently always takes second place. The latter seems to be a comparatively 
rare insect in Lagos. Apart from those bred from larvae, a few adults were cap- 
tured in crab-holes in the Golf-course swamp in July 1914, and occasional specimens 
have been obtained in Government Quarters in the residential area. The larval 
sources in the period 1914 to 1918 were as follows :—Wells, 6 occasions ; boat 
or canoe, 2; barrel, 1; total, 9. If a slight but constant degree of salinity is 
favourable to this species one would expect a greater prevalence. O. wrritans is 
a very common mosquito in Lagos and probably prefers a breeding-place with 
an appreciable amount of chlorides. It seems to have a special proclivity towards 
crab-holes, but an equally common source is surface pools ; it also occurs in wells, 
catch-pits, and various receptacles of domestic or public utility ; roof-guttering, 
canoes, and banana-tree stumps have yielded occasional samples, Curiously enough 
O. nigricephalus, its common associate in crab-holes, does not appear to affect 
domestic receptacles, not even wells, yet the adult insect is commonly captured 
in houses. 

Culex decens, which freely uses crab-holes as a breeding-place, is after S. fasciata 
the most common species in wells ; other sources were surface pools, drains, pots, 
tubs, etc., and once each a tree-hole, agbo pot, and rain-water on a tarpaulin. 

Culex tigripes, from the predaceous habit of its larvae, is rather a welcome assistant 
than otherwise, though its presence deranges one’s estimate of other domestic 
species. It has been found in wells, pools, pots, and once in agbo. 

Culex duttont seems to be less common in Lagos town than at Ebute Metta on the 
mainland. Water-pots and wells, and once a canoe, were the only sources discovered, 
but it has been previously found at Yaba in dirty water in tubs, barrels and ponds. 

Culex fatigans (see above). 

Culex grahami has been bred from well samples, from surface pools and borrow- 
pits, and once from a water-barrel. 

Stegomyra luteocephala has its chief habitat in trees. The only domestic source has 
been the roof-gutters (from which it has also been obtained in Accra) ; a crab-hole, 
a pool in a yard, and a telegraph-pole socket have furnished larvae on isolated 
occasions. 

Culex quartt was obtained on two occasions from water-pots ; it has previously 
been found in a pond at Yaba. 

Culex insignis occurred once in a yard well; crab-holes and surface pools are 
usual sources. 

Culex ataenvatus was obtained once only, from a household pot. 

Culex thalassius was obtained once from a water-bucket ; away from dwelling- 
houses its sources are numerous, viz., pools and brackish swamps, crab-holes, canoes, 
and once it was found in a railway girder. 

S. africana was hatched on one occasion from a sample from a household tin ; 
it has also been obtained from surface pools, and previously from a bamboo at Yaba. 

Eretmopodites quinquevittatus was found abundantly on one occasion in a 
potsherd in a garden in the Hospital Compound. 


266 J. M. DALZIEL. 


Anopheles costalis takes a surprisingly high place amongst those obtained from 
household sources, especially as pools within the compound fence are excluded. In 
Lagos it can be reckoned upon to supply from 2 to 3 per cent. of the total finds 
in house to house inspection and generally takes third place numerically. Graham 
(loc. cit.), finding the average reaching 6°3 per cent. in February, a dry month, 
suggests that the reduction of puddles out of doors tends to a relative increase 
of Anopheles larvae in domestic receptacles. The present observations, however, 
are against this, as the finds of A. costalis here recorded occurred chiefly in the wet 
months. Thus out of 74 occasions, 52 occurred in the months June to September, 
and 64 in the months May to October, inclusive. In the year 1916, when no samples 
were examined in the months May to October, this species is entirely absent from 
the list. In the Accra list quoted above, A. costalis similarly takes third place 
amongst those obtained from domestic sources. Of domestic samples, wells con- 
tribute nearly half the number for this species in Lagos, viz., 35 out of 74. The 
water-pot filled at the well may thus receive the larvae, but there is little doubt 
that direct use is made of pots, tins, etc., within the house. On two occasions the 
larvae were found in agbo pots, and on six, in catch-pits ; drain, roof-gutter, banana- 
tree stump, and even a coconut shell furnished occasional samples. In the present 
series none occurred in tanks and barrels which receive rain-water from roofs. 

The larvae of A. mauritianus were found once only in the roof-gutter of a 
Kuropean’s residence. 

The electric cat-fish (Malapterurus electricus) is not infrequently kept in 
domestic water-pots by the people of Lagos, its presence being supposed to preserve 
the purity of the water. Although it tends to act as a deterrent to the sanitary 
inspector, and therefore to neglect of cleansing the utensils, it certainly feeds on 
mosquito larvae, as was proved by experiment at the Research Institute at Yaba. 


Collections of Adult Mosquites. 

The native sanitary inspector, whose task is the thorough and regular inspection 
of a daily block of compounds, is not a suitable person to rely on for collection of 
mosquitos in native houses, unless this duty is allotted to a definite mdividual 
relieved of other duties for the time. Although considerable numbers have been 
obtained and identified at one time or another, they have usually been collected 
either by the medical officer of health himself or by the European sanitary inspector, 
and few of these collections have been representative of the native town. Relative 
frequency of different species 1s greatly affected by locality and time of year. Thus 
in some low-lying streets in Lagos town a collection taken in the rains would 
certainly contain an overwhelming proportion of A. costalis. On one occasion 
a collection made by a sanitary student resident in Ebute Metta (on the mainland) 
consisted almost entirely of Mansonioides africanus, which is a rarity in Lagos 
town proper. 

Attempts were occasionally made to interest the members of the class of sanitary 
students in training to collect adult mosquitos in the premises where they lodged, 
and cyanide tubes were issued for the purpose. The results were below expectation 
and the youths soon tired of the good work. The following, however, gives the 
result of one such effort representing native houses in different parts of the native town. 


CRAB-HOLES, TREES, AND OTHER MOSQUITO SOURCES IN LAGOS. 267 


(As the main object of the collection was to ascertain the relative prevalence of 
S. fasciata, Anophelines are excluded, their numbers being always excessive during 
the period covered; two specimens of A. nili occurred amongst several 
hundreds of A. costalvs). 


TasBLe VIII. 
Adult Culicine Mosquitos in Native Houses, Lagos, July to September 1917. 


| 
Species. No. of Specimens obtained. | Percentage. 
Culiciomyia nebulosa | 83 74°8 
Ochlerotatus nigricephalus | 10 9°0 
Culex decens ie om: et od 5 4°5 
> grahami i as a 3 yA | 
Ochlerotatus irritans .. - ee! | 3 2:7 
Stegomyia fasciata .. ed ees 3 td 
Culex tigripes .. ; | 2 1:8 
» thalassius 2 1:8 
Total | 111 


This list might be regarded as corroborating the remarks above on the influence 
of the agbo pot, and of the sanitary inspector, on the preponderance of Culicoomyia 
over Stegomyia, were it not that Ingram and Macfie warn us against accepting such 
a collection (probably made chiefly in the evening) as evidence of the relative 
absence of S. fasciata, a scarcity which is often demonstrable even in blocks where 
its larvae are predominant, and for which a possible explanation is that this species 
“ whilst entering houses to obtain a feed of blood, does not roost indoors ” (Bull. 
Ent. Res. vii, p. 173). More abundant collections under proper conditions of 
observation should settle this point, but material of this nature from Lagos is at 
present insufficient. There seems no doubt that under native social conditions 
in Lagos C. nebulosa has become a highly domesticated species, and this habit 
along with its actual abundance suggests the need for a study of its bionomics 
and potentialities as a possible carrier of disease. 

Many mosquitos have been captured for identification at various times in the 
Kuropean quarters to the east of the Race-course and around the Golf-course. The 
following list (p. 268) shows one such classified collection, in which note was taken 
of the numbers obtained during one month only; in a total of 1,682 specimens 
a rich mosquito fauna is represented including 26 species. 

June is the month of greatest rainfall and A. costalis preponderates. Much 
has been done since then in the area in question in the way of filling hollows and 
draining. ‘The abundance of crab-holes in the locality no doubt influences the order 
of precedence amongst the Culicines. O. domesticus takes a comparatively high 
place, though its larvae have only once been discovered in a neighbouring swamp ; 
they have previously been found in borrow-pits by the railway on the mainland. 

A smaller collection of 150 mosquitos from the same houses, covering the period 
15th May to 21st July, 1917, is shown overleaf; 16 species are represented. 


J. M. DALZIEL. 


TABLE IX, 


Adult Mosquitos captured in European Quarters, June 1914. 


Species. No. me eto Percentage of Culicines. 
Ochlerotatus irritans 112 26:23 
s nigricephalus 70 16°39 
Culiciomyia nebulosa 53 12-41 
Culex thalassius .. # 45 10°54 
Ochlerotatus domesticus .. 25 5°85 
Culex decens 23 5°38 
Stegomyia fasciata 13 3°04 
Mansonioides africanus TO * 2°34 
Culex consimilis .. 9 2°11 
Ochlerotatus cumminsi 8 1°88 
Culex fatigans 7 1:64 
Uranotaenia annulata 7 1°64 
Culex grahami 6 1-40 
» ligrupes .. a 5 1°16 
Banksinella punctocostalis 5 1:16 
Culex duttoni ae 4 "93 
» quasigelidus 2 °47 
Ochlerotatus caliginosus es 2 °47 
> argenteopunctatus .. 1 23 
a punctothoracis 1 23 
Culex salisburiensis ‘ 1 "23 
Taemorhynchus annettt .. 1 23 
% aurites .. a 1 23 
Others undetermined .. ae 16 3°75 
Total Culicines .. 427 
Anopheles costalis se oe 1251 Culicines. 25°4% 
re mi .. dis a 3 Anophelines 74°6 % 
9 mauritianus .. ats 1 
Total Anophelines 1,255 
Total Mosquitos 1,682 
TABLE X. 


Species. 
Ochlerotatus irritans .. st. 
% migricephalus 


Culiciomyia nebulosa . . 
Oulex decens 
Anopheles costalis 
Ochlerotatus domesticus 
Culex grahami : 

» insignis 

3 TING  .. ay 
Taeniorhynchus aurite 
Culex thalassius 

» aduttont .. 

» pruina .. 
Stegomyia fasciata 

: luteocephala 

Anopheles nili S 


Total 


No. of Specimens obtained. 


ee DO OO HP IR CO 


150 


Percentage. 


mem DD 09 


Pi igh: Agsindh. Aye 
woWIASAIASS 


each 0°7 


CRAB-HOLES, TREES, AND OTHER MOSQUITO SOURCES IN LAGOS. 269 


The following species obtained in this locality on other occasions may be added 
to the above lists, viz.:—Hodgesia sanguinis, Theo., O. ochraceus, Theo., C. unwittatus, 
Theo., Mansonioides uniformis, Theo., Taeniorhynchus metallicus, Theo., Urano- 
taenva balfouri, Theo., U. bilineata, Theo., var. frasert, Theo. The last-named was 
at one period fairly abundant in the neighbourhood of Government House, but the 
others are of rare occurrence. 

On 31st August 1915, a strong tornado from N.E. occurred about 6 p.m., in the 
course of which very numerous mosquitos were blown into the quarters occupied 
by the Medical Officer of Health. An examination of 36 specimens captured for 
identification gave the following curious result :—Culex decens, 339, 13; Culex 
rma, 12; M. africanus, 19. 

There is here a lack of correspondence with the record of mosquitos taken in the 
same rooms during the immediately preceding 3 months, which is as follows :— 
403 mosquitos comprising A. costalis, 375 or 93 per cent.; O. nigricephalus, 8 ; 
C. nebulosa, 6; C. thalassius, 3; C. decens, 2; O. writans, 2; O. domesticus, 2; 
S. fasciata, 2; C. quasigelidus, 1; M. africanus, 1; T. metallicus, 1. 

It is probable that on this occasion some particular nidus close at hand was 
stirred up. A search further off in the Ikoyi Plains to windward of the populated 
area, where there are swamps and small surface pools, crab-holes, etc., as well as 
a few out-lying hamlets, revealed chiefly A. costalis, O. wrritans, U. annulata, and 
M. africanus. 

The numerous breeding places on the island beyond the limits of the inhabited 
town, and again the mangrove and brackish swamps beyond the island itself across 
the creeks to south and east, are doubtless capable of furnishing Lagos, at least 
intermittently, with wind-blown mosquitos, and possibly some species rarely 
found as latvae in the local sources may have their origin in these regions beyond. 

In the period dealt with in these notes which may be taken as from May 1914 
to August 1918, the number of different species which have come to notice in Lagos 
Municipal Area has been 46,7 viz.:— 


ANOPHELINAE. 
Anopheles costalis, Theo., A. funestus, Giles, A. mauritianus, Grandpré, 
A. obscurus, Griinb.,* A. nili, Theo.* 


CULICINAE. 

Banksinella punctocostalis, Theo.* 

Culex ataenratus, Theo., C. consimilis, Newst., C. decens, Theo., C. duttoni, Theo., 
C. fatigans, Wied., C. grahami, Theo., C. guiarti, Blanch., C. imsignis, Carter, 
C. pruina, Theo.,* C. quasigelidus, Theo.,* C. rima, Theo., C. salisburiensis, Theo., 
C. sitiens, Wied., C. thalassius, Theo., C. tgripes, Grandpré, C. univittatus, Theo.* 


t To these 46 may be added 10 species absent from this list but included amongst the 
41 species recorded from Lagos and published in the pamphlet of the Yellow Fever 
(West Africa) Commission, ‘ Distribution of Mosquitoes in West Africa,” Aug. 1913, 
viz. Anopheles pharoénsis, Theo., Aédomyia africana, N.L., Banksinella lineatopennis, 
Ludl., Hretmopodites inornatus, Newst., Mimomyia mimomyiaformis, Newst., Ochlero- 
tatus longipalpis, Grinb., O. marshalli, Theo., Stegomyia apicoargentea, Theo., Toxorhyn- 
chites brevipalpis, Theo., Uranotaenia mashonaensis, Theo. 


(713) F 


270 J. M. DALZIEL. 


Culiciomyia nebulosa, Theo. 

Eretmopodites quinquevittatus, Theo. 

Hodgesia sanguinis, Theo.* 

Micraédes inconspicuosus, Theo. 

Mansonioides africanus, Theo., M. uniformis, Theo.* 

Ochlerotatus apicoannulatus, Edw., O. argenteopunctatus, Theo.,* O. caligi- 
nosus, Grah.,* O. cumminsi, Theo.,* O. domesticus, Theo., O. irritans, Theo., 
O. nigricephalus, Theo., O. ochraceus, Theo.,* O. punctothoracis, Theo.* 

Stegomyia africana, Theo., S. fasciata, F., S. luteocephala, Newst. 

Taenorhynchus annettr, Theo.,* T. aurites, Theo.,* T. metallicus, Theo.* 

Uranotaenra annulata, Theo., U. balfourt, Theo.,* U. bilineata, Theo. var. frasert. 

(N.B.—Species marked with an asterisk have not been found in the larval state 
within the period dealt with in this paper, but the larvae of C. quasigelidus, 
O. caliginosus, O. punctothoracis, and U. balfoury have been recorded by Graham 
from borrow-pits at Yaba (Bull. Ent. Res. 1, pp. 32, 39, 50). Breeding-places of 
the other species are referred to in the text.) 

Of the 5 species of Anopheles, A. costalis is universally distributed over the island 
and is very abundant in the rains ; it also makes an appreciable figure in the returns 
from domestic receptacles. Its eradication is one of the chief objectives of sanitary 
effort. 

A, mauritianus is a rarity, but it has been bred from larvae in a roof-gutter 
and captured once or twice in European quarters ; the larvae have also been taken 
in a pond and in road puddles at Yaba on the mainland. 

A. funestus was bred from slightly brackish water in a canoe. 

A. obscurus was captured once in a native house, but has not been taken in the 
larval state. 

A. nili is found now and then in European houses in the Golf-course and Race- 
course residential area, and in offices and native houses in the town. Its larvae 
have not been obtained, but no doubt have their origin in the stagnant waters 
which abound on the island. The imagines when found have always been in such 
very scanty numbers that its influence as a malaria carrier seems almost negligible 
at present. The circumstances which favour the prevalence of one species over 
another are, however, incompletely understood, and as these may conceivably 
alter in unexpected ways or in conditions beyond immediate control, it is well to bear 
its presence in mind, and to discover and deal with its sources. For similar reasons 
the above notes and record of species found in Lagos within recent years, even 
if little more than a bare recital of names, may at any time prove of interest and 
possibly of practical use to future workers. 


SOME NEW INJURIOUS WEEVILS. 
By Guy A. K. Marswatt, C.M.G., D.Sc. 
(Puate VIL.) 


Subfamily BRACHYDERINAE. 


Tanymecus destructor, sp. nov. (Pl. vii, fig. 5). 
Colour piceous black ; the head with dark brown scaling above, except for a 


line of pale scales bordering the eye, and with pale buff scales beneath ; scaling 
of the prothorax brown or grey brown on the disk, an ill-defined dark brown 
stripe at the side, and grey or buff beneath ; scaling on the elytra grey or brown, 
with a very ill-defined darker stripe between striae 2 and 5, usually vanishing 
at a short distance from the base, but often traceable to behind the middle in the 
form of spots. 

32. Head with shallow, confluent and longitudinally striolate punctation ; 
the forehead at its narrowest not narrower than the base of the rostrum, and without 
any central fovea or carina ; the eyes very short oval, moderately convex. Rostrum 
a little shorter than the head, gradually narrowed from base to apex, and the dorsal 
area more markedly so, so that both scrobes are visible at the same time directly 
from above for almost the whole length; the dorsal area flat, sculptured like the 
head, and with a mere trace of a carina on the anterior half, the apical margin only 
shallowly simuate. Antennae with the scape reaching the hind margin of the eye ; 
the funicle with joint 1 longer than 2, and 2 as long as 3+4; 3to7 subequal. Pro- 
thorax a little longer than broad (9) or a little broader than long (9), broadest 
at the middle, the sides rather strongly and regularly rounded ; the apex and base 
of equal width and both slightly arcuate, the apical margin rather oblique in lateral 
view ; the dorsum evenly convex and with fine confluent punctation throughout. 
Scutellum inconspicuous. LElytra subtruncate at the base, separately rounded 
at the apex, with very sloping shoulders and without any humeral angle, the sides 
gently rounded and markedly narrower in the than in the Q, and without any 
posterior callus; the striae fine, shallow and shallowly punctate; the intervals 
even, almost flat and finely shagreened ; the numerous short recumbent scale-like 
setae hardly distinguishable from the narrow true scales. Legs densely clothed 
with variegated scales and setae ; the front femora thicker than the others, the 
two anterior pairs bearing long erect hairs beneath on the basal half inthe ¢; the 
hind tarsi with joint 1 as long as 2+3, and 2 longer than 3. 

Length, 6-8 mm.; breadth, 2-3 mm. 

S. Ruopesia: Salisbury, 5,000 feet, xi-xii. 1894 (G.A.4.2.); Mazoe, 4,000 
feet, xii. 1919 (R. W. Jack). 

Described from 17 3 g and 19 9 9Q. 

Of the described South African species, this insect is most nearly allied to 
T. inafectatus, Fhs., which has similarly sloping shoulders, the scape reaching to the 
hind margin of the eye, and similar hind tarsi. On the other hand the latter is a 
larger insect (10-12 mm.), with the rostrum as long as the head; joint 2 of the 

(713) F2 


pi" a GUY A. K. MARSHALL. 


funicle is as long as 1, and longer than 2+3; the scales on the elytra are much 
smaller, etc. 

This wingless terrestrial species is abundant in the Salisbury district, and accor- 
ding to Mr. Jack, Government Entomologist, Southern Rhodesia, the adults some- 
times cause considerable damage to the maize crops by eating off the young plants 


as soon as they appear above ground, They also feed on the leaves of other crops, 
such as sweet potatoes. — 


Tanymecus agricola. sp. nov. (Pl. vu, fig. 6). 


Colour black, variegated with greyish buff scaling and light and dark brown 
setae; the pronotum with three indistinct dark stripes; the elytra with several 
indefinite interrupted dark lines. 

$2. Head with rather coarse longitudinally confluent punctation ; the forehead 
not narrower than the base of the rostrum and without any central fovea or carina ; 
the eyes almost circular and rather strongly convex. Rostrum as long as the head 
and parallel-sided, the apex asymmetrically emarginate in the middle, the nght 
side of the angle being rounded and the left straight; the dorsal area parallel- 
sided, shallowly impressed in the middle, with a well-marked narrow median 
carina, the sculpturing like that of the head. Antennae with the scape 
elongate, reaching beyond the front margin of the prothorax; the funicle 
with the two basal joints equal, 2 equal to 3+4, and 3 to 7 subequal. Prothorax 
longer than broad in both sexes, gently rounded at the sides, broadest a little before 
the middle, and with a faint apical constriction; the apex slightly narrower than 
the base, and both margins very gently arcuate, the apical margin oblique at the 
sides; the dorsum evenly convex, with close confluent punctation throughout ; 
the short broad recumbent setae pale on the areas of pale scaling and dark brown 
elsewhere. Scutellwm elongate, with dense whitish scales. Hlytra jointly sinuate 
at the base, with rounded humeral angles, parallel-sided from the shoulders to 
beyond the middle, and impressed on each side before the apex, which is jointly 
rounded ; the striae rather deep, especially towards the apex, but partly hidden 
by thescaling towards the sides, the punctures large at the base and becoming smaller 
behind ; the intervals not broader than the punctures on the disk, slightly convex 
and shagreened ; the pale areas formed of short oval or almost circular scales, 
mixed with numerous pale short flat curved setae, the dark areas only with similar 
dark brown setae. Legs grey, the femora with an indistinct pale spot beyond 
the middle; the front femora slightly thicker than the others, the two anterior 
pairs in the g clothed with long hairs beneath on the basal half; the hind tarsi 
with jomt 1 as long as 2 + 3, and 2 equal to 3. 

Length, 7-8 mm. ; breadth, 2:°2-2°8 mm. 

S. Rnopesta: Gwelo, i. 1920 (R. Lowe Thompson). 

Described from 2 3g and 2 99. 

Although very similar in general facies to 7’. destructor, this species may readily 


be distinguished by its longer and parallel-sided rostrum, more convex eyes, much 
longer scape, broad scales, etc. 


It has also been found feeding upon maize. 


SOME NEW INJURIOUS WEEVILS. 273 


Subfamily OTIORRHYNCHINAE, 
isaniris ater, sp. nov. (Pl. vii, fig. 1). 

Colour uniform dull black, the elytra with a small spot formed of small elongate 
white scales near the base of the 5th row of punctures, and sometimes smaller ones 
on the 7th and 9th, also a lateral row of similar spots near the margin, and 
occasionally traces of another a little above it. 

$9. Head with coarse shallow punctation and a deep median furrow, and sparsely 
set with short spatulate white setae; the eyes moderately convex, deepest in the 
middle. Rostrum broad, gradually dilated at the apex, which is deeply sinuate in the 
middle, the epistome not carinate on its posterior edge; the dorsal area broad, 
regularly widening to the apex, broadly impressed in front, almost flat in the basal 
half, and there coarsely punctate and with a broad smooth median carina, which 
is highest at the base; the basal incision gently curved; a longitudinal lateral 
ridge running from the scrobe to the eye. Antennae black, with recumbent pale 
setae ; the scape not much curved, moderately slender and rather abruptly clavate ; 
the funicle with joints 1 to 6 progressively diminishing, 7 as long as 4. Prothorax 
broader than long, strongly rounded at the sides, broadest at the middle, truncate 
at base and apex, the base being narrowly carimate and a little broader than the 
apex ; the dorsum covered with large flattened contiguous granules mostly forming 
irregular pentagons, each with a setiferous puncture ; laterally the granules become 
elongate and less defined. Hlytra narrowly ovate in the 3, much broader in the Q, 
broadest not far from the base, each with about 18 rows of punctures running in 
pairs, but becoming rather irregular towards the sides and apex; the intervals 
between the pairs more regular than the intervening ones, no broader than the 
septa in the rows and becoming granulate on the apical third, each bearing a spaced 
row of short subrecumbent setae, which are longer and more erect on the apical 
area. Legs rather stout, clothed with coarse pale setae and sparse setiform white 
scales, the setae being much longer, denser and softer on the lower edge of all the 
tibiae in the g; the hind tibiae of the ¢ deeply excised on the inner edge for more 
than half their length from the apex, the excision flattened and smooth. 

Length, 8-8°75 mm. ; breadth, 3:2-4°2 mm. 

S. Ruopesia: Salisbury, ix. 1893, ete. (G.A.K.M.—type). NYASALAND: 
Blantyre, iv—v. 1910 (Dr. J. E. S. Old). 

The sculpturing of the pronotum will distinguish this insect from all the described 
species of [saniris, except I. acuticollis, Fst., which differs in having the prothorax 
sharply angulate behind on each side and in lacking the granules on the elytra. 
In general form and structure it much more resembles such species of Systates as 
S. brevicollis, Fst., and S. crenatipennis, Frm., but these differ in having only 10 
rows of punctures on the elytra, which Faust considered to be the only character 
separating Systates from Isanirvs. 

This weevil occurs very commonly on trees of the genus Brachystegia, which 
form the principal component of the woodland on the Mashonaland plateau ; 
but Mr. R. W. Jack informs me that it is a very general feeder, attacking various 
cultivated plants, including citrus trees. 


Systates exaptus, sp. nov. (PI. vii, fig. 2). 

Colour black, with brownish grey “scaling, which is sufficiently thin on the pro- 
notum and disk of the elytra for the shiny integument to show through, but is 
denser at the sides and especially on the declivity of the elytra ; above the declivity 
an ill-defined common transverse dark chevron-shaped patch, the posterior edge 
of which is deeply sinuated. 


274 GUY A. K. MARSHALL. 


32. Head obsoletely punctate, the forehead with a short deep median furrow ; 
the furrow separating it from the rostrum almost straight ; the eyes strongly convex, 
deepest in the middle. Rostrum as long as broad ( Q) or slightly longer than broad 
(3), subparallel-sided in its basal half, then gradually dilated to the apex; the 
dorsal area alutaceous, shallowly impressed on each side of the sharp median carina, 
the lateral margins with shallow confluent punctures, very slightly costate, parallel 
in the basal half, then gradually diverging; the central carina broadly forked a 
little beyond the middle, the fork enclosing a large smooth subtriangular impressed 
area just behind the epistome; a low longitudinal ridge running from the scrobe. 
to the eye. Antennae with the scape slender, cylindrical, abruptly clavate and 
clothed with recumbent white setae and narrow setiform scales; formula of funicle: 
1, 2, (3, 7), (4, 5, 6). Prothorax broader than long, gently rounded at the sides, 
broadest at the middle; the apex truncate or shallowly sinuate, a little narrower 
than the base, which is feebly marginate ; the dorsum closely set throughout with 
low contiguous shiny granules, each bearing a puncture containing an erect scale- 
like seta; laterally the granules are much reduced and interspersed towards the 
base with coarse shallow punctures. Elytra without any truncate margin at the 
base, very broadly ovate in the 9, much narrower in the 4, widest not far from the 
base, the shoulders broadly rounded, the posterior declivity steep in the § and almost 
perpendicular in the 2; the striae rather shallow, closely punctate, the punctures. 
as broad as (9) or narrower than (Q) the intervals, each of the latter bearing a 
single row of short erect flattened setae; the scales small, narrowly ovate, the 
palest ones broader. Legs with short pale setae and narrow scales; all the tibiae 
denticulate internally in both sexes, the hind pair in the ¢ broader, with the inner 
face flattened and smooth throughout and incurved at the apex, the lower edge. 
deeply bisinuate and with an obtuse tooth a little before the middle; in the 92 
the hind tibia has a short smooth shallow hollow on the inner face near the apex. 
Venter of 3 with two widely separated small tubercles near the posterior edge of 
the Ist segment. 

Length, 6°5-7°5 mm. ; breadth, 3-3°6 mm. 

S. Ruopesia: Salisbury. i. 1895—i. 1905 (G.A.K.M.—type); Unmtali, xi. 
1901 (G.4.K.M.); Mpudzi R., Umtali dist., xi. 1901 (G.4.K.M.); Mt. Chirinda, 
Melsetter dist., i. 1907 (C. F. M. Swynnerton). 

This common species appears to feed on a number of different herbaceous plants, 
and the adults have been observed by Mr. R. W. Jack to do appreciable injury 
to young maize plants,* 


* The following closely allied species occurs on the coast in the same latitude :— 
Systates beiranus, sp. n.—Closely resembling S. ewaptus in form and colouring, but 
smaller and differing in the following particulars : rostrum with the dorsal area slightly 
convex longitudinally in the basal half, the anterior impressed portion not triangular, 
but separated off by a low transverse ridge ; no longitudinal ridge behind the scrobe ; 
the epistome slightly asymmetrical, the apex of the curve formed by the delimiting 
carina being a little to the left of the median line, and the carina on the right side rather 
longer and higher than that on the left; the prothorax with a very low median costa 
in the anterior half ; the hind tibiae of 3 smooth and flattened internaily, but less 
incurved at the apex and without any tooth on the lower edge ; the elytra vertically 
truncate at the base; Ist ventrite without tubercles inj}. Length, 5-6 mm. ; breadth, 
aa mm. PortTuGUESE East AFRica: Beira, x.1900—x.1904 (P. A. Sheppard) ; 

S327 QO 


bo 
~I 
Ol 


SOME NEW INJURIOUS WEEVILS. 


Systates chirindensis, sp. nov. 


32. Dull black, uniformly and thinly clothed with very small grey or light brown 
scales, sometimes with a greenish reflection ; in the ° only there is a large common 
triangular denuded dark patch at the top of the declivity with its apex directed 
backwards. 


Head separated from the rostrum by a straight furrow, impunctate, and with a 
shallow median furrow that is longer than half the transverse one; the eyes 
moderately convex, deepest in the middle. Rostrum a little broader than long, 
very slightly narrowed from the base to the middle, then gradually widening to 
the apex ; the dorsal area almost flat, finely alutaceous, parallel-sided to the middle 
and then dilated, the margins slightly costate and very shallowly punctate, the 
median carina low and broadly forked beyond the middle, the fork enclosing a 
shallowly impressed subtriangular smooth bare area; no costa behind the scrobe. 
Antennae long and very slender, clothed with sparse fine recumbent setae; the 
scape cylindrical, abruptly clavate ; the relative lengths of the joints of the funicle 
thus: 1, 2, 3 (4, 5), (6, 7). Prothorax somewhat broader than long, moderately 
rounded at the sides, broadest a little before the middle, truncate at base and 
apex, the former not marginate and a little broader than the apex; the dorsum 
evenly set with low shiny granules, each with a setigerous puncture ; laterally the 
granules are much reduced. Slytra very narrowly ovate in the g, much broader 
in the 9, without any truncate margin at the base, the apices jointly rounded and 
produced downwards like a beak, especially in the 9; the striae shallow and 
containing punctures that are deep towards the base but diminish much behind ; 
the intervals broader than the punctures, slightly convex, each with a row of small 
low setigerous granules, the setae being longer and thinner in the Q, and also duphi- 
cated on the declivity of interval 1 in that sex. Legs thinly clothed with narrow 
pale scales, the hind femora with a broad dark band in the middle; in the 4, the 
front tibiae slightly bent inwards at two-thirds from the base and denticulate 
internally, the mid pair also denticulate and the lower edge very strongly curved 
in the apical half, the hind pair only slightly curved, the inner face broadly flattened 
and smooth almost to the base, with a single row of granules along its lower edge 
in the apical half and the upper edge granulate ; in the Q, the tibiae only slightly 
curved, the hind pair with a smooth flat area on the inner face bounded above 
and below with a ‘row of granules. Venter with ventrite 1 in the ¢ very broadly 
impressed at the base and with two widely separated small conical tubercles on the 
hind margin. 


Length, 64-8 mm. ; breadth, 2°6-3°6 mm. 


S. Raopes1a: Mount Chirinda, 4,000 ft., xu. 1901 and x. 1905 (G.A.K.d1L). 
Described from 14 gg and 15 99. 


From the very few species of Systates having granulated elytra, such as S. granosus, 
Fst., this insect may be distinguished by the absence of the vertically truncate edge 
at the base of the elytra. 


In 1915, Mr. C. F. M. Swynnerton found the adults of this species feeding in 
numbers on the leaves of coffee bushes on Mt. Chirinda. 


276 GUY A. K. MARSHALL. 


Subfamily CRyPTORRHYNCHINAE. 
Genus Elytroteinus, nov. nom. 


Pteroporus, Fairmaire (nec Schonh. 1843), Ann. Soc. Ent. France, 1881, p. 307. 

To the generic characters given by Fairmaire the following may be added :— 
The mesosternum parallel-sided, much broader than one of the mid-coxae, the 
posterior edge of the cavity markedly overhanging ; the intercoxal process of the 
metasternum very broadly truncate, no metepisternal suture; ventrite 2 in the 
middle much longer than 3 + 4; the elytra with stria 10 complete, and the lateral 
margin rather strongly smuate at the base; the front femora with a stout tooth, 
and all the femora very shallowly sulcate beneath; the tibiae with a sharp carina 
on the dorsal edge ; the tarsal claws simple and divaricate. 


Elytroteinus subtruncatus, Frm. (Pl. vii, fig. 8). 

This is the only known species of the genus and was originally described from 
Fiji (op. cit. p. 308), but it has recently been found by Mr. O. H. Swezey at Honolulu 
attacking the roots of ginger (Hedychium coronarvum). 


Subfamily CALANDRINAE. 


Calandra shoreae, sp. nov. (Pl. vii, fig. 4). 

3 2. Colour dull black or piceous, without scaling ; the rostrum, legs and antennae 
red-brown. 

Head with three transverse rows of punctures behind the eyes, the forehead 
with an elongate fovea. Rostrum elongate, slightly curved, gradually widened 
at the apex, with four dorsal rows of punctures and three lateral ones on each side, 
of which the uppermost disappears in the basal half; in the 9 the rostrum is very 
little longer than that of the g and the punctures are finer. Antennae with joints 
3 to 5 of the funicle about as long as broad, the others longer. Prothoraz a little 
longer than broad, widest near the base, the sides slightly rounded and gradually 
narrowed to the apical constriction; the base slightly rounded in front of the 
scutellum and truncate on each side, the apex broadly sinuate; the dorsum 
coarsely and closely punctate, the transverse septa between the punctures being 
narrower and lower than the longitudinal ones, producing the effect of numerous 
undulating longitudinal carinae; the median carina well-marked, fairly straight, 
and running from the apica! constriction almost to the base; each puncture with 
a very short stout erect pale seta. Hlytra with the usual sculpturing characteristic 
of the genus, intervals 1, 3, 5, 7 and 9 being rather higher and much broader than 
the others, and each with a row of punctures almost as large as the quadrate ones 
in the striae ; the posterior callus feeble, and intervals 3 and 9 not elevated at the 
apex. Abdomen with the exposed part of the pygidium forming almost an 
equilateral triangle, flat and coarsely scrobiculate. 

Length, 3°2-4'4 mm. ; breadth, 1:2-1°8 mm. 

Inp1a: Dehra Dun, United Provinces (C. F. C. Beeson); Kheri, U.P. (Beeson) ; 
Gorakpur, U.P. (A. E. Osmaston); Shwegu, Burma (Beeson); Pusa, Bengal ; 
Adoni, Bellary dist., Madras; Pattikonda, Kurnul, Madras; Waltair, Madras 
(S. W. Kemp); Nilgiri Hills (WH. ZL. Andrewes). Mauritius (J. £. M. Brown). 


SOME NEW INJURIOUS WEEVILS. 277 


Very closely allied to Calandra sculpturata, Gyl., from South and East Africa,* 
which has the pronotum somewhat similarly sculptured, but the longitudinal cari- 
nation is not so well marked in the basal half and the median carina is less distinct 
in the African species ; this also has a small spot composed of a few pale scales in 
the middle of the base of the pronotum, and another on the shoulder of each 
elytron ; and the exposed part of the pygidium is more elongate and transversely 
convex. C. shoreae attacks the seeds of the sal tree (Shorea robusta), as well as 
those of Dipterocarpus turbinatus. 

Prof. A. Hustache had already drawn up an M.S. description of this species upon 
two specimens from Mauritius, but he has kindly suggested that the present 
description should be published instead. 


Calandra glandium, sp. nov. (Pl. vu, fig. 3). 

Closely allied to the foregoing species, and differing principally in the following 
characters :—A rather larger insect and longer in proportion to its breadth ; rostrum 
of the 9 finely aciculate, rather dull, and as strongly punctate as that of the ¢ of 
C. shoreae; the rostrum of the g much shorter and straighter, rugosely punctate 
and with four dorsal rows of minute granules; the pronotum with large reticulate 
punctures, all the interspaces being equally raised and the median carina usually 
indefinite and abbreviated ; the elytra longer in proportion, the posterior callus 
more developed and intervals 3 and 9 elevated at the apex, so that the apical 
junction of striae 3 and 8 lies in a deep impression ; the rugose area of the pygidium 
more elongate and with a sharp median longitudinal carina. 

Length, 4°6-5'2 mm. ; breadth, 1°6—-1°8 mm. 

Inp1a: W. Almora, Kumaon, numerous specimens bred from acorns of Quercus 
ancana (H. G. Champion—type) ; Naini Tal, Kumaon, bred from acorns of Quercus 
dilatata (H. Marsden). 


Stenommatus musae, sp. nov. (Pl. vii, fig. 7). 

Colour dull piceous black, usually with a greyish sheen ; the antennae, legs and 
apical portion of the rostrum red-brown. 

Head short globular, smooth, rather shiny, with minute sparse punctures ; the 
eyes actually contiguous beneath. Rostrum () elongate, as long as the head and 
prothorax together, slightly curved, moderately stout, almost cylindrical but some- 
what flattened beneath, not dilated at the apex, opaque and with small close shallow 
punctures for four-fifths of its length, the apical fifth impunctate and very shiny; 
a deep lanceolate median furrow from the base to the antennae ; the lower surface 
impunctate, with a low shiny median carina; colour red-brown, with only the 
basal third or fourth part blackish ; in the 3, a little shorter and broader pro- 
portionately, and blackish from the base to the middle or beyond. Antennae with 

* Schénherr records this species from Bengal, but his description agrees well with a 
South African species. Prof. Aurivillius has kindly confirmed the identification on 
comparing one of my African specimens with the type of C. sculptwrata, which, he 
informs me, bears the label ‘‘ Cap. B. spei ? Drége.”” The Indian record seems almost 


certainly erroneous. Drége resided in South Africa, and all the other species described 
from his collection by Schonherr are from that country. 


278 GUY A. K. MARSHALL. 


the scape short, just reaching the eye. Prothorax a little longer than broad, the. 
sides subparallel from the base to beyond the middle, then roundly narrowed and 
very deeply constricted near the apex, the constriction continued across the dorsum 
as a deep curved furrow ; the apical margin truncate or very slightly sinuate, the 
base truncate ; the dorsum gently convex transversely and evenly set throughout 
with small separated punctures, the interspaces being as broad as the punctures. 
themselves ; on the sides the punctures are rather larger and much closer together. 
Elytra elongate-ovate, slightly sinuate jomtly at the base, and without any hu- 
meral prominence ; the sides gently curved, broadest at a short distance from 
the base, then rather rapidly narrowing to the apex, which is broadly rounded ;. 
the striae containing rows of shallow distant punctures, which encroach slightly 
on the sides of the convex impunctate intervals. 

Length, 1°6-1'8 mm.; breadth, 0°6-0'7 mm. 

Hawalan Is.: 2 ¢ 3, 8 2 Q, Kaimuki, Oahu, 19.11.1920 (0. H. Swezey). 

The genotype, S. fryz, Woll., from Mexico, differs in its slightly larger size ; the. 
scape extends behind the eye ; the eyes are not quite confluent beneath; the head 
is opaque and coarsely punctate ; the rostrum is markedly shorter ; the prothorax. 
is broader and much more coarsely punctate, the interspaces being much narrower 
than the punctures, etc. 

This species was found by Mr. Swezey breeding in numbers in the root (corm): 
of a banana plant. 


ame me 


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wy ico 

> . as" 
ped 


Fee ’ 
: at 
: - 7 : > 7 
4 ‘2 1) e 
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eG. | 8 ; 
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Fig. 


EXPLANATION OF PLATE VII. 


. Lsamris ater, sp.'n., c- 
. Systates exaptus, sp. D., . 
. Calandra glandium, sp. n., &. 


‘ shoreae, sp. n., &. 
Tanymecus destructor, sp. n., . 
ms agricola, sp. D., 3. 


. Stenommatus musae, sp. Nn. 
. Elytroteinus subtruncatus, Frm. 


PraTe VIf. 


PART 8. 


XI. 


VOL, 


Butt. ENT. RESEARCH. 


INJURIOUS CURCULIONIDA. 


279 


THE HABITS OF A DIPTERON PREDACEOUS ON MOSQUITOS 
IN NYASALAND. 


By W. A. Lamporn, M.R.CS., L.R.C.P. 


On 12th September 1919, while watching at a pool near Kotakota, Lake Nyasa, 
the emergence of mosquitos from their pupae, which in the case of this particular 
species takes place mostly between 4.30 p.m. and dusk (about 6.30 at that time 
of year), the writer saw at rest on some scum on the surface of the water, an An- 
thomyiid fly of the genus Lispa, which after a few minutes made a sudden rush 
at a mosquito just emerging from its pupa-case. The mosquito with its wings 
still unexpanded and part of its legs and the hinder segments of its abdomen still 
within the pupa-case was absolutely at its mercy. The fly grappling it from behind 
with its front legs proceeded to thrust its proboscis into the thorax. The fly was 
by no means timid, and it was possible to lean over and approach one’s head to watch 
the proceedings so closely that 1t was even possible to determine by its antennae 
the sex of the unlucky mosquito; it was a male. The fly was captured before 
it had finished its meal, lest it should suddenly fly up and escape. 

Next morning the pool was again visited and quite a number of the same flies 
were observed on the scum. One was seen to make a sudden thrust with its pro- 
boscis into the water, at the edge of the scum, followed by a series of similar but 
smaller efforts, as if it had seized some body, though insecurely at first. Presently 
it raised itself and partly drew out of the water a mosquito larva, about half-grown, 
impaled on its proboscis, which it proceeded to suck. The larva was not seen to 
move, possibly being paralysed by some buccal secretion from the fly. Advantage 
was taken of the fly’s lack of timidity to push beneath the scum a broad stick 
on which it and its victim were gradually raised, so that the feeding process could 
be studied as near at hand as one wished. A good series of the flies, subsequently 
taken, were all removed in this manner, the stick being thrust into the mouth of a net 
which was then closed. 

One fly was seen on the same occasion to grasp and thrust its proboscis into a 
mosquito, a female, which had failed to emerge completely from its pupa and may 
or may not have been dead at the time; it was not seen to move at all. The 
rest of the series were all seen at one time or another attacking mosquito larvae, 
not always with success, but when successful dragging them out of the water and 
sucking their juices. 

On 13th September, the emergence of a Culex was witnessed almost from the 
first ; the thorax bursting its covering, the imago then extracting its head, the rest 
of the body following until the wings and some of the legs were free. At this point 
a Lispa, which had been resting about four inches off on some scum, approached 
deliberately on the surface film, grasped the still soft Culex with its forelegs, and 
thrust its proboscis into the thorax, dragging the mosquito beneath its body to a 
little piece of leaf, about two inches off, floating on the water. As the captor 
walked, one could see quite clearly the legs of the victim trailing beneath. The 
fly with its prey was then taken. The mosquito, a female, was still alive and 
moved its legs feebly. 


280 W. A. LAMBORN. 


The attack of another Lispa in a similar way on a mosquito pupa was witnessed 
on the same occasion. The pupa impaled on the fly’s proboscis, was drawn from the 
water, fully extended, to a little bit of leaf on which the fly had been sitting. It 
was not seen to move, so that one is unable to assert that it was living when seized 


by the fly. 


At a later date many instances were witnessed of attacks by the flies on mos- 
quitos which had been unable to fly away on the previous night owing to the failure 
of their wings to expand properly. One fly had impaled in the usual way a helpless 
mosquito, and had dragged it on to a small stick in the water. On an attempt 
being made to secure it, the insect flew across the water bearing its prey to another 
support, a foot further off in the water. Threatened by another attempt it flew 
up bearing its prey and settled fully another foot away on a blade of grass where 
both were captured. Its prey, a female mosquito, moved freely after capture. 
The manoeuvres of another fly were precisely the same when it was menaced with 
the net. In view of the unsuspicious attitude of most of the flies previously cap- 
tured, this wariness was a little surprising, more especially as they were taken, 
with the water still in the shade, at an early hour when most insects of diurnal 
habits are prone to be sluggish in their perceptive faculties. 


The pressure of other work made it impossible to study the behaviour of at least 
four other Diptera, all having business on the water. One species appeared 
to act as a scavenger, endeavouring to extract nourishment out of empty pupa- 
cases and larval skins of mosquitos. Another small species, which was not captured, 
was almost certainly predaceous on mosquito larvae. It gave from time to time 
most careful and deliberate attention to the cleaning of its long, pointed proboscis, 
aiter which it gradually stalked the smaller mosquito larvae breathing at the surface 
film, endeavouring, seemingly, to thrust its proboscis into their air-tubes, and 
repeating this action again and again, though in no instance was a successful issue 
witnessed. 


In the same pool in which these Culex larvae occurred in swarms were a few 
predaceous larvae of a much larger species of mosquito, which every now and again 
attempted to secure one of the Culex larvae, causing them instantly to scatter. 
On the far side of the pool, which was about two feet across, was noticed a “ rat- 
tailed ” larva struggling on the surface, and on near approach it was seen that one 
of the larger species of mosquito larvae, almost fully grown, had seized it firmly 
at the point of junction of the “ tail” with the body. The rat-tailed larva wriggled 
and contorted itself in every possible way; its tail became twisted round the 
head of its adversary at least twice; and it endeavoured to escape by diving into 
the depths of the pool, but unsuccessfully. The mosquito larva never slackened 
hold and, when its victim approached the surface, immediately put up its respiratory 
siphon and probably nibbled away constantly, for by and by a portion of the cuticle 
of the larva was seen to be partly detached near the mouth of its assailant and 
to be trailing the water. Towards the end of a prolonged and desperate struggle 
a second similar mosquito larva joined in the attack, seizing the Syrphid larva at 
a point midway between the two extremities of its body. The Syrphid struggled 
still more violently. Its new assailant, better situated for attack at a vital spot 


HABITS OF A DIPTERON PREDACEOUS ON MOSQUITOS. 281 


than the original one, very soon bit a hole through which more and more of the 
Syrphid’s viscera gradually protruded. Eventually this mosquito larva dropped 
off and wriggled away, taking a small portion with it, much of which was consumed 
at leisure. The Syrphid’s struggles thereafter got more and more feeble and in 
two or three minutes it straightened out and ceased to move, floating on the suriace, 
the first mosquito larva still holding to it. An attempt was then made to secure 
this larva, which however alarmed by the movements of the numerous Culex larvae 
through the water being disturbed by the net, suddenly let go and vanished. The 
precise species of larva is therefore impossible of determination ; all that can be 
recorded in referrence to it is that 1t was considerably larger than any of the Culex 
larvae; it had posterior respiratory spiracles only; it also suspended itself in 
the water rather more obliquely than the Culex larvae did.* It was not possible 
to capture any with the means at one’s disposal. 

Such a fate for these Syrphid larvae would seem to be by no means uncommon, 
for on the following morning two more, both dead and with viscera protruding, 
were seen floating on the surface. One rather wondered how it was that any of the 
Syrphid larvae, which were securely grouped on a piece of rotting cassava one 
and a half inches below with their tubes thrust up to the surface, ever came up 
to the top, thereby incurring such risks. The explanation is probably to be found 
in the occasional failure of the larvae to withdraw their life-lines from the surface 
in sufficient time to escape the sudden attack of the mosquito larvae and in their 
subsequent endeavours to set themselves free. 

When the sun was off the pool full-fed Syrphid larvae, with their tails already 
partly shrivelled, were seen to crawl out and to burrow in dry sand well above 
waterline, where their puparia were readily obtained. Some of these larvae placed 
on 12th September in a box containing dry sand pupated in about forty-eight hours, 
and the flies emerged on about 23rd September, the pupal stage having therefore 
occupied about twelve days. The species has been identified by the Imperial 
Bureau of Entomology as Phytomia curta, Lw. During the heat of the day these 
flies were frequently seen ovipositing in the wet sand at the margin of the pool. 

[The fact that Lispa preys on mosquito larvae has also been recorded by 
Mr. J. Mitford Atkinson (Jl. Trop. Med. xii, 1909, pp. 255-6 ; Howard, Dyar & Knab, 
Mosquitoes of N. America, i, 1912, p. 170), who in Hong Kong observed Lispa sinensis, 
Schiner, devouring larvae almost as long as their own bodies.—Ed.] 


* [The predaceous larvae were probably either Toxorhynchites brevipalpis, Theo., or 
Culex tigripes, Grp., both of which are known to prey on other mosquito larvae.—ED. } 


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283 
A NEW TYPE OF ENTOMOLOGICAL KILLING-BOTTLE. 
By Matcotm E. MacGrecor, 
Wellcome Field Laboratory, Surrey, Wellcome Bureau of Scientific Research. 


The “cyanide bottle” is never a very attractive part of the entomologist’s 
equipment. When made up in the old way, 7.e., plaster of Paris poured over lumps 
of solid potassium cyanide and the plaster then allowed to set, the contrivance 
has an attractive appearance, but is an abomination to use, especially if small 
insects are being dealt with. The large amount of water utilised in making up the 
plaster of Paris can never be dried out properly, and the potassium cyanide being 
deliquescent to some extent the presence of moisture is unavoidable. Moreover 
the cyanide soon decomposes and turns brown, imparting its colour to the plaster 
and forming an ugly-looking mass on which small insects are not contrasted and 
may be easily overlooked and lost. 

The newer method of making up the killing-bottle by powdering the cyanide 
and mixing this with about an equal weight of powdered borax, compressing the 
mass at the bottom of the bottle, and afterwards pouring over it a small amount 
of plaster just sufficient to secure the powders, is a distinct improvement in so far 
as the approach to anhydrous conditions is concerned and the more rapid liberation 
of HCN gas. However, the mass decomposes more rapidly with the addition 
of the borax, and the unfortunate brown coloration is accentuated. 

There are too the methods that depend on fixing the potassium cyanide in the 
cork instead of in the bottle, which are certainly improvements in that the insects 
are left lying on the glass, against which they can be easily seen; but with this 
method the cyanide is not in a very safe situation, and furthermore cannot so 
rapidly evolve as much HCN gas as cyanide that is distributed over the bottom 
of the bottle. 

In addition to the faults peculiar to whatever method is employed in making 
up the cyanide killing-bottle there are the following disadvantages in the use of 
cyanide :—(1) the bottle is a potential danger to the ignorant if not properly looked 
after, owing to the highly poisonous nature of potassium cyanide ; (2) if the bottle 
is accidentally broken, cases of poisoning may result unless every scrap of the 
cyanide and plaster is collected and destroyed; (3) the killing power of the 
preparation speedily weakens if the bottle is constantly used ; (4) insects recover 
more often from HCN gas in weak concentration than they do from concentrated 
chloroform vapour; (5) when the cyanide bottle is exhausted it is preferable to 
throw the whole thing away (unless the bottle is of the type where the cyanide is 
secured in the cork) rather than to attempt to chip out the hard plaster to renew 
the apparatus, and a new killing bottle has therefore to be obtained. 

On the strength of these objections I have for several years preferred to use 
chloroform as an insect poison, utilising chloroform-saturated rubber in place of 
cyanide. The method is ideal as far as its action is concerned, but the one great 
drawback has been the rapid evaporation of the chloroform and the constant 
addition of the fluid that has been necessary. 


284 MALCOLM E. MACGREGOR. 


In order to have a killing-bottle which will have the great advantage of chloroform 
as the poison, the bottle described below is now used. Jt can be made from 
ordinary laboratory material, unless a very large-mouthed bottle is required. 

Select an ordinary “ boiling-tube ” (large form of test-tube), and a large rubber 
cork which will just slide down the inside of the tube, making rather a tight fit. 
Bore a hole in the centre of the cork so that a small piece of glass-tubing (internal 
diameter, 2 to 3 mm.) can be forced through. The glass-tubing should be of such 
a size that it projects about # inch on one side of the cork, and about + inch on the 
other. 

Now slide the cork into the boiling-tube so that the ? inch length of glass tubing 
projects towards the bottom of the boiling-tube, and force it down until the end 
of the tubing touches the bottom of the tube. If the rubber cork is a tight fit 


roinary 
cork. 


-- Glass 
bolling-tube. 


_ Za’ projection 
of glass tubing. 


on - Red rubbercork. 


s %’, ojection 
Chleroform.~ of glass tubing. 


Fig. 1. A chloroform killing-bottle. 


(as it should be) moisten the cork slightly with water and press it down the boiling- 
tube by the aid of the open end of a small test-tube. Warm the apparatus so as 
to drive off any moisture that may remain, and when dry it may be charged with 
chloroform. This is accomplished as follows :—Pour into the boiling-tube suff- 
cient chloroform to stand about 2 inches above the upper surface of the rubber 
cork, and then carefully warm the air-space below the cork over a spirit-lamp. 
The air will be partly driven out by expansion. Remove the tube from the flame 
and as the space cools the chloroform will run in to fill the partial vacuum. Then 
again heat the bottom of the tube until the chloroform within boils gently, and all 
the air is driven out by the chloroform vapour. On cooling once more the space 
will this time be completely filled with chloroform. Now invert the tube and 


A NEW TYPE OF ENTOMOLOGICAL KILLING-BOTTLE, 285 


pour out the surplus chloroform from the upper surface of the cork. None of the 
chloroform in the space below will escape, owing to the air pressure at the upper 
open end of the glass tubing. Gently warm the sides of the tube above the cork 
to drive off any remnants of liquid chloroform, and when an ordinary cork has 
been fitted, we have an apparatus similar to that shown in fig. 1. 

Although pure rubber is soluble in chloroform, it will be found that the compound 
from which the rubber corks (red variety) are made, swells when in contact with 
chloroform but will not dissolve. 

The following are the advantages of the apparatus :—(1) A high concentration 
of chloroform vapour is always available; (2) if the tube is accidentally broken 
there is no danger of poisoning ; (3) 5 cc. of chloroform is sufficient for a week’s 
continuous use, and the tube is therefore exceedingly economical. 

Care should, however, be taken that the lower end of the tube containing the 
chloroform be not unduly heated by the hand, otherwise gaseous pressure will 
force the liquid up the central glass tubing. Should this occur at any time it is only 
necessary to shake out the few drops of liquid at the upper end of the cork, and 
blow into the tube to dry the cork surface. If the tube is inverted, no amount 
of overheating will do more harm than to dissipate the chloroform to some extent, 
and for this reason it should be carried in this position. The open lower end of 
the glass tubing then stands above the surface of the liquid chloroform, and the 
apparatus is “ sealed.” 


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Be ae 
beak’ tre h) Mc A 


287 


INSECT PESTS OF DATES AND THE DATE PALM IN. MESOPOTAMIA 
AND ELSEWHERE. 


By P. A. Buxton, M.A., F.E.S., 


Fellow of Trinity College, Cambridge ; formerly Entomologist, 
Mesopotamian Expeditionary Force. 
(Pirate VIII). 


In the summer of 1918, there was every prospect of an unusually good crop of 
dates in Mesopotamia, until the month of June, when a large number of the half- 
grown dates suddenly became “ hashaf’’—an Arabic word signifying that they - 
dried up and fell from the trees. This was a very serious matter, not only because 
the date is the principal resource of the country, but also because any and every 
disaster was at that time attributed by a section of the population to the British 
administration. At the end of July I was lent to the Directorate of Agriculture 
to carry out an investigation upon the failure of the date crop. I found that the 
principal pest, the larva of a Gelechid moth, had finished its work, and had wan- 
dered away from the palms to pupate ; and though I was unable to solve the prob- 
lem of its life-history, I had the good fortune to discover a good deal that was new 
about the insect pests of the date palm. At the time I published a report (1918), 
to which I only refer now in order to say that it was produced on the spot and without 
access to any books ; it contains certain inaccuracies, and anything of value which 
I may have discovered will be found in the present paper. During the course of 
my investigations I visited all the important date-growing areas of Mesopotamia, 
that is to say, Baghdad and its neighbourhood, the Diyala River from Ba’qubah 
to the Persian Frontier at Baba Pillawi, Mendali and Balad Ruz, Basra and the 
country round it, Mohammerah and Fao, the Lower Euphrates from Nasiriyeh 
to Sugq-ash-Shuyukh, Qurnah and Amara. At the time of my visit the fruit was 
nearly ripe, so that it was not difficult to learn the distinctions between the different 
varieties of date palms, a point of great importance in view of the fact that they 
are liable to the various pests in varying degrees. I strongly recommend future 
investigators to familiarize themselves thoroughly with these varieties, and also 
with the very considerable number of Arabic words which are used specially for 
the date palm and its cultivation. This is no small task, as will be realized when 
I mention that in Mesopotamia about 70 words are employed by the peasants for 
various parts of the date palm ; the vocabulary relating to cultivation and methods 
of employment is just as extensive. 

The literature relating to the pests of the date is so scattered, some of it in the 
most obscure periodicals, that I believe this paper will meet a want. My aim 
in writing it has been to deal with the pests observed in Meopotamia, but to give 
at the same time a summary of all that has been written up till the present time, 
and a full bibliography. From the bibliography I have excluded all purely popular 
references to well-known pests, all reports, etc., which merely record the presence 
of one of these pests in some place in which it is well known to occur, and papers 
relating solely to the systematics of the pests. 

(713) G2 


288 P. A. BUXTON. 


I am indebted to Prof. R. Newstead, Mr. E. E. Green, Mr. G. J. Arrow and 
Mr. J. H. Durrant for much help most kindly given, and to the staff of the Imperial 
Bureau of Entomology, who have always assisted me in every possible way. 


LEPIDOPTERA. 
Family HESPERIIDAE. 
Padraona palmarum, Moore. 
This butterfly is recorded by Stebbing from leaves of date palms at Calcutta. 


Family PYRALIDAE. 
Subfamily PHycirinaE (= ANERASTIINAE). 
Myelois phoenicis, Durrant. 

This species was described by Durrant (1915) from a specimen bred by Lord 
Walsingham in 1904 at Hammam-es-Salahin, Constantin, Algeria. The larva was 
found in dates. The species has smce been found in England and France in im- 
ported dates, and as Durrant remarks, “may be widely spread by commerce.” 


Plodia interpunctella, Hb. 

According to Popenoe the Indian meal moth is a serious pest of stored dates 
in Arizona. This is the only record I have found of this omnivorous and very widely 
distributed species attacking dates. 


Spermatophora hornigii, Léderer. 

Moths which I bred at Basra in September 1917, and Baghdad in September 
1918, from larvae found in August feeding in wind-fall dates, have been identified 
by Lord Rothschild as this species. He informs me that Léderer’s original speci- 
mens also came from dates, but I have been unable to consult the original reference. 
The species appeared to be quite common in Mesopotamia, but was in no sense 
& pest. . 


Ephestia calidella, Guenée. 

Gough (1917) mentions that in Egypt the larvae of this species sometimes attack 
living dates on the tree, and bore into them through the rind. These larvae spin 
sulk at all times in their life-history, and lower themselves on a silken thread when 
they are alarmed. In these respects the habits of this insect differ from those 
of the Ephestia in the Khargeh oasis. This species occurs at Baghdad and Amara, 
but I have no evidence that it is a pest. 


Ephestia cautella, Walk. (Hphestia passulella, Barrett.) 

This species is a common warehouse pest in Basra, and probably throughout 
Mesopotamia. The larvae also enter windfall dates in the gardens in Baghdad, 
Amara, and Basra, passing into the fruit at the poimt at which it has become 
detached from the perianth. These windfall dates are frequently harvested by the 
less well-to-do, who in this way infect and reinfect their stores and godowns with 
E. cautella. It is a common thing to see masses of dates which have been riddled 
by this and other Pyralid moths exposed for sale in the bazaars ; these spoilt dates 
are bought for feeding domestic animals. 


INSECT PESTS OF DATES IN MESOPOTAMIA. 289 


The larva of Z. cautella not only eats ripe dates which have fallen to the ground, 
but also the small “ hashaf’’ dates which have become hard and withered owing 
to the attacks of the Gelechiid moth (see below) while they were half grown. 

According to Smyth the life-history from egg to adult occupies five weeks in 
summer at Smyrna; in April, May and June about 48 days. Chittenden says 
that this species has a wide distribution in Europe, Asia, North Africa and North 
and South America; it was originally described from Ceylon. 

It has been recorded as attacking pomegranates, pears, cotton cake, cacao beans, 
gall-nuts, flax-seed meal, tonka beans, walnuts, pecan nuts, pea-nuts, figs, pearl 
hominy, dried moths (specimens), stored corn, asparagus berries, white rice, 
chick pea, zizyphus berries, etc. 

Chittenden and Smyth’s Bulletin should be consulted for details of the life-history 
of this pest, and for an account of the damage it does to stored figs. Buckler 
described its life-history (under the name Hphestia passulella) in 1882. 

It is parasitised in Egypt by Habrobracon kitchenert, Dudgeon & Gough. 


Ephestia sp. ? 

Gough (1917) describes the life-history of an Hphestia which is not identified but 
differs from £. cautella and E. calidella. This larva is a pest of fallen and harvested 
dates in the Dakhlah and Khargeh oases, which lie in the desert west of the Nile. The 
. life-history is given as follows. The female lays as many as 190 eggs in 24 hours, 
and this is not the full number; the eggs are deposited im ones and twos on the 
surface of the dates after they have fallen or been gathered. The larvae wander 
about and are not able to enter living dates, a point in which they differ from the 
larva of H. calidella. They generally enter a stored date by the calyx end, and 
take up their abode in the space between the flesh and the stone. After feeding 
for a minimum of 20 days, during which no silk is spun, the larvae emerge from the 
date and wander about, leaving a silk thread behind them; they wander for two 
days, leaving their thread on all the dates over which they pass, and then spin a 
cocoon in some convenient corner. The minimum life period from egg to adult 
is 25 days in August, 27 in September, and 38 in October, in Egypt. The species 
passes the winter in the larval stage. The adult moths are crepuscular and 
nocturnal, and never fly far. 

Gough suggests that arrangements should be made for marketing the dates 
in two grades, and that the first grade should be treated with SO, after they are 
packed ; the second grade should be pressed, a process in which many of the larvae 
are crushed. He also speaks of the employment of moth-proof receptacles. I 
imagine that it will be some years before the inhabitants of these desert oases use 
either moth-proof receptacles or sulphur fumigation, but they can probably be 
induced to gather up their wind-falls and give them to their cattle to eat. 


Subfamily GALLERIINAE. 


Arenipses sabella, Hampson. 
The larva of this species is known to eat stored dates, to which so far as we know 
it is confined ; in this respect it differs from E'phestia spp., which are pests of the 


290 P. A. BUXTON. 


most catholic tastes, devouring almost all forms of stored produce. The larva is 
described by Durrant as dull pinkish-brown, head and plates darker, brownish 
spots darker, length 1 im. This species has been recorded from Fao, at the head of 
the Persian Gulf, in Arabia, also from Algeria and Persia (Hampson). As an 
imported species it has reached England, and has been reported from Canterbury 
by Hampson; in the British Museum there is a specimen bred from the larva 
described by Durrant; this larva was found by Miss B. Reed in dates in London. 
The species was common at Amara on the R. Tigris, males and females coming 
to ight from Aprilto June. It seems probable that Mesopotamia is one of the native 
countries of this sect. There is no record of it as a serious pest, but this may 
only be due to our ignorance of the moths whose larvae attack stored fruits. 


Family GELECHIIDAE. 


Genus and species unknown (text fig. 1). 

This was by far the most serious pest of the date palms in Mesopotamia in 1918, 
and it was on account of its ravages that I was ordered to report on the failure of 
the date crop during that season. Unfortunately I received this order after the 
larvae had left the dates to pupate, and it is only owing to a fortunate chance 
that I was able to obtain a few larvae which had been preserved in whisky by one 
of the officers of the Department of Agriculture. These specimens enable me 
to say that the pest is a Tineid of the family GELEcHIIDAE, if the tables given by 
Fracker for determining Lepidopterous larvae are correct. I have found his paper 
most useful, and I believe it to be accurate, but I think we must accept it with a 
certain amount of reservation, at any rate where the Microlepidoptera are concerned, 
because Fracker has been forced to base his conclusions on the examination of a 
comparatively small number of identified species of larvae. Text fig. 1 shows the 
chaetotaxy of this larva, and with the aid of it future investigators should be able 
to determine whether any pest with which they are dealing is identical with the one 
of which I am speaking. The explanation of the nomenclature of the setae will 
also be found in Fracker’s paper. Apart from chaetotaxy the larva may be 
described as follows. The adfrontals reach the vertical triangle, and the frontals 
about half way up to the vertical triangle. The superior angle of the frontals is 
not prolonged into a fine point. The arrangement of the ocelli is that characteristic 
of the GELECHIIDAE and not the OECOPHORIDAE, a point which cannot be described 
without reference to a figure (cf. Fracker, Pl. vit, figs. 80, 81). The body is 
slightly flattened, and is widest at the third and fourth abdominal segments ; the 
setae in general are short and inconspicuous, as is usual in internal feeders. The 
prothoracic shield is weakly chitinised and is not darker than the rest of the body ; 
the legs are short. The abdominal prolegs are also short, and the crochets on 
them are uniordinal, and arranged in a complete series: the crochets of the anal 
prolegs (text fig. 1,D) are arranged in a continuous transverserow. The prothoracic 
spiracle is oval, the abdominal ones circular, that on the 8th segment the largest. 
[ was informed that the larvae when alive were pink, without any markings. 

This pest has apparently not been recorded before, and is so far as we know 
confined to Mesopotamia. Its biology is very little known, but ‘its ravages were 
first detected by other observers in June. The half-ripe dates turned brown and 


INSECT PESTS OF DATES IN MESOPOTAMIA. 291 


became spherical, and inside them the larva could always be found eating the flesh 
of the date along the stone; early in July the larva left the dates, which were 
by now quite dry and about half the size of a healthy date, making its egress by 
a hole bored close to the attachment of the perianth. From the hole one could 
always see frass and silk projecting. It is extremely rare to see two emergence 
holes from one date, and I believe that the eggs must be laid singly, probably on the 
fruit soon after it is set, and that the larva burrows into it and remains in a single 
date throughout its period of growth. All my efforts to find the place of pupation 


Fig. 1. Structural details of larva (Fam. Gelechiidae) causing damage to half-ripe 
dates in Mesopotamia. A. Chaetotaxy of pro and meso-thorax (I, II) and 3rd, 7th 
and 8th abdominal segments (3, 7, 8). The shield of the prothorax is drawn rather 
too wide and too deep. The head of the larva is supposed to be towards the left. 
C. Left proleg of 7th abdominal segment seen from below; the seta ¢ is in the 
direction of the mid-ventral line. D. Anal proleg seen from above. 

Small Greek letters (a, 8, 7, w, ete.) indicate individual setae, according to 
Fracker’s terminology. Greek capitals (B, K, etc.) indicate groups of setae, of 
regular occurrence in certain families or many families of the Lepidoptera, e.g., 
B=a-+8, P (capital rho)=e+tp, T=r+w+¢ (2), K=n-+x. Or., crochets of proleg; 
leg, proleg, or leg; sp., spiracle ; sh, dorsal shield of prothorax. 


were unsuccessful. I searched cracks on the palm stem, the fibrous leaf-sheaths 
and the rubbish which accumulates in the crown of the tree; I also passed earth 
irom beneath the trees through a sieve, without finding any pupae except those 
of large Nocrurpaz. I believe that pupation takes place in the ground, and lasts 
throughout the winter, because we always observed that the incidence of the pest 
was slight in gardens in which a crop of winter and spring vegetables was cultivated 
beneath the trees. I opened a very large number of the dates which had fallen 
prematurely as a result of the attack of these larvae, and found no dead larvae, 
so it appears that the pest is not controlled by parasitic Hymenoptera or 
entomophthorous fungi. 


292 P. A. BUXTON. 


This pest attacks all varieties of date palm, but it was particularly harmful to 
the Khustawi in Baghdad and Ba’qubah, and the Sair (Ista’mran) and Halawi 
in the Basra neighbourhood. The age of the tree appeared to make no difference 
to the severity of the attack, but it was an invariable rule that a heavy loss was 
sustained by owners of gardens far from the river, and by those who failed to keep 
the ground beneath the trees clear of weeds. Palms growing in rather salt ground, 
and those irrigated by the rise and fall of the actual sea at Fao were not particularly 
attacked by this insect. It will be understood that only a few sorts of palm will 
live under these conditions, but those that lived did not appear to be any less healthy 
than individuals of the same sort irrigated by fresh water. I have already said 
that the pest was never very serious in gardens in which green vegetables were 
cultivated beneath the trees ; I believe that this was due to the breaking up of the 
soil, and consequent damage to pupae, but we cannot regard this as proved until 
we know more of the biology of the insect. It was suggested that the very heavy 
incidence of this pest in the year 1918 was due to the fact that many of the cul- 
tivators had taken work in the Labour Corps, with the result that the gardens 
were less dug and ploughed than usual. 

Until the life-history of this very serious pest is known, one cannot do better 
than impress on land-owners and cultivators the need to dig the ground beneath 
the trees in winter ; but if it is found that the female moth oviposits on the flowers 
of the female palm, or on the young fruit soon after it is formed, it might be worth 
while to protect the whole inflorescence with a muslin bag. Popenoe figures such 
bags in use to defend the ripening dates from wasps and birds. 

It is impossible to estimate the loss due to this larva in 1918; I found evidence 
of its ravages in every garden I visited from Baghdad to Fao, and from Nasiriyeh 
on the Lower Euphrates to Khaniqin on the Kurdish border. Its predilection 
for the Sair dates of Basra is serious, as this is the most widely grown of all sorts 
and its fruit is exported to India in immense quantities. The more costly sorts 
suffer least, because no man owns more than a few trees, and these are always well- 
watered and carefully cultivated ; I refer to such sorts as Barhe, Braim, Maktum, 
and Mirhaji. I made a rough estimate that the following percentages of the crop 
of all varieties were lost, between the time when the fruit set and the harvest ; 
at Ba’qubah 50-70 per cent., at Qurnah 40-50 per cent., at Basra 30-40 per cent. 
The Basra date forest is the largest in the world, and contains about 7,000,000 
palms. One may assume that the average yield of a palm is 60 lb. of fruit, and 
it will be realized that the loss of about a third of the crop is an extremely serious 
matter. This is the pest which I wrongly referred to in my report (1918) as a 
Pyralid. The attacked dates are called “ hashaf” in Arabic. 


COLEOPTERA. 
Family DYNASTIDAE. 


Oryctes elegans, Prell. 

In Mesopotamia this species, for the identification of which I am indebted to 
Mr. G. J. Arrow, is common throughout the date-growing area: I have seen speci- 
mens from Baghdad, Ba’qubah, Balad Ruz, Mendali, Amara, Basra and Fao. 


INSECT PESTS OF DATES IN MESOPOTAMIA. 293 


Adults fly to light from April to September and do great damage by boring through 
leaf-bases and stems of inflorescences in the crown of the tree. I have seen a 
garden littered with bunches of nearly ripe dates felled by this insect, which can 
be a most serious pest. I presume that the females lay their eggs in the burrows 
they make. Beetles of this species may also frequently be found in burrows in the 
hard trunks of living or dead palms, and I believe that these burrows are made 
either by the adult or the larva, but I have no proof that this is so. When these 
burrows are numerous they interfere seriously with the nutrition of the tree, and 
they sometimes cause it to break off in a high wind. The larva of a large Lamelli- 
corn, which I believe is this species, lives in the crown of the date- palm among 
the leaf-sheaths, etc., and bores its way through the bases of leaves, the bases 
of fruit-bearing stems and the young leaves before they unfold. Sometimes it bur- 
rows inwards towards the actual growing point. It appears to have a predilection 
for fairly young trees, between the 10th and 20th year. It is an interesting fact 
that the larva of O. rhinoceros occasionally adopts the same habit, and lives in the 
crown of coconut trees, especially if they are decayed (Burkill); it is of course 
well-known that this larva normally inhabits piles of vegetable refuse, and old 
thatch and palm leaves which are lying on the ground. Such a habit would I 
think be impossible for the larva of O. elegans ; for owing to the extreme desiccation 
of Mesopotamia it is generally impossible to find any decaying material of any sort 
in the gardens. 


Treatment.—The presence of the adult or larva in the crown of a palm is known 
from the immense quantity of chewed vegetable fibre which passes through them 
and falls from the mouth of the burrow. When he sees this the Arab cultivator 
climbs the tree and endeavours to extract the insect with the short heavy knife 
with which he prunes his palms, and in so doing he frequently commits much 
havoc himself. 


As Prell’s O. elegans is very little known and was described from the female 
I have asked Mr. Arrow to re-describe the species. His description which is as 
follows is founded on a series of ten collected by W. E. Evans, C. F. C. Beeson 
and others. 


“ Oryctes elegans, Prell. This species has hitherto been known from its female 
only, examples of that sex taken at Fao having been described in a German perio- 
dical in July 1914 (Entom. Mitt. in, p. 210.) It is one of the smallest repre- 
sentatives of the genus Oryctes, males ranging from 28 to 34 mm. in length, females 
from 34 to 36 mm. It is narrower in shape and rather less convex than O. deser- 
torum, Arrow (found in the same region) and the upper surface is more smooth 
and shining, the elytra being almost devoid of the punctation occurring on the 
latter species, but possessing a deeply incised line on each side of the sutural edge 
instead of the row of dots found in O. desertorum. . The two points of the clypeus 
diverge more widely and are separated by a rounded, instead of an angular excision. 
The armature of the head and prothorax scarcely differs from that of the other 
species, but the posterior edge of the thoracic cavity is rather more sharply toothed 
in the female, and rather less so in the male, which has only a very slight blunt 
projection there, not distinctly bicuspid as in O. desertorum. The cephalic horn 


294 P. A. BUXTON. ‘ wi 


in the male is short, but in well-developed specimens longer than in the female. 
The thoracic cavity is similar in both sexes, except for the sharper posterior tooth 
in the female just mentioned. The pygidium is recurved in both sexes and has 
a fringe of red hairs on its free edge only; in the male it is convex and 
very smooth and shining; in the female it is more strongly inflexed and rather 
strongly and closely punctured. The last ventral segment in the male is also very 
smooth and its front and hind margins are straight and parallel; in the female 
it is strongly punctured, and forms a rounded lobe.”’ 


Oryctes desertorum, Arr. 
Oryctes desertorum, G. J. Arrow, Fauna Brit. India, Coleoptera Lamellicornia 
(Cetoniimae and Dynastinae) 1910, p. 276, 

The distribution of O. desertorum is given by Arrow as “ India, Karachi; Persia; 
Arabia, Masqat (Muskat), Lahej and Fao.” He tells me that the Persian specimen 
came from South Persia, so that all the records are from countries in which the 
date-palm is grown, It appears probable that O. desertorum will eventually prove 
to be a date-palm pest; at present however we know nothing of its habits or life- 
history. 


Oryctes rhinoceros, L. 

Burkill gives Phoemx dactylifera in a list of palms attacked by “ the Rhinoceros 
Beetle.” Stebbing records O. rhinoceros from date palms at Madras. On the 
other hand Copeland in his very careful account of the pests of the coconut does 
not mention the date palm in the list of trees which O. rhinoceros is known to attack, 
nor does Ghosh. O. rhinoceros is found from India eastwards, so that its range 
and that of the date palm only coincide in parts of India. It may well be that 
in that country it attacks this palm, as Stebbing has stated, but I do not feel that 
the matter is proved in view of the fact that the species of Oryctes have until recently 
been much confused. The life-history of O. rhinoceros is well-known, as it is a very 
serious pest of coconut. Ghosh’s and Burkill’s papers and Copeland’s book should 
be consulted for details about this member of the genus and the measures which 
have been adopted to destroy it. 


Family NITIDULIDAE. 
Carpophilus dimidiatus, F. 
Popenoe speaks of this species (as ‘‘ Carpophyllus dimidiatus”’) eating stored dates. 


The ordinary “ Dried Fruit Beetle” is Carpophilus hemipterus, L, The parts of 


Popenoe’s book which deal with pests of the date tree are not as valuable as the 
rest of the book. | 


Family ANOBIDAE. 
Coccotrypes ? dactyliperda, F’. 

Willcocks records small beetles, which he believes to be this species, from Amry 
and Aglawy dates from the Sharkia province of Egypt. As many as 11°8 per cent. 
of the latter variety have been found infected. The beetles live ina cavity hollowed 
out of the date stone, in which as many as forty eggs, larvae, pupae and imagines 
may be found. A small circular hole communicates between the cavity and the 
exterior. 


‘ 
ia 


INSECT PESTS OF DATES IN MESOPOTAMIA. 295 


Family CURCULIONIDAE. 


Rhynchophorus ferrugineus, Oliv. 

Popenoe states that the “‘ Coconut Palm Borer, Rhynchophorus ferrugineus, has 
killed date palms in India, and to less extent in Mesopotamia.’’ Opposite p. 168 
he gives a plate of two dying palms nearly leafless, under the title “ Ravages of 
the Palm Borer.’’ However in a footnote he says that “ he did not see the borer 
itself in Baghdad,” and his record appears to rest on descriptions received from 
Arab sources. As far as I am aware this very large and conspicuous weevil has 
never been obtained in Mesopotamia. Captain W. Edgar Evans, R.A.M.C., never 
saw it though he lived in Amara, an important date-growing centre, for about a 
year and made a very extensive collection of beetles; it is absent also from one 
or two smaller collections of beetles received from that country by the British 
Museum. 

In India, on the other hand, this weevil is definitely known to attack the date 
palm, and MacKenna tells us that it is capable of inflicting considerable damage, 
It is found that the palm may be saved from attack if mud enclosures are 
built round the trunk and kept filled with water. Milne figures palms the tops 
of which have bent right down owing to this weevil. It is important to cover 
with earth or tar the raw surface from which an offset has been detached. This 
weevil is distributed from India to the Philippines (and possibly to New Guinea 
also). Burkill gives a list of half a dozen palms from which it has been recorded, 
the chief of them being the coconut, of which it is a major pest. 

It will possibly be found that other species of Rhynchophorus, e.g., the African 
f. phoemcis, occasionally attack date palms. 


THYSANOPTERA. 
Family THRIPIDAE. 


Heliothrips haemorrhoidalis, Bch. 

This insect is recorded by Williams as having been found on date palm in 
Trinidad. It is the species known as the ‘“‘ Greenhouse thrips,’ and has been 
recorded from every continent and has a long list of food-plants. It is not a pest of 
the date palm. 


RHYNCHOTA. 
Family Coccipae. 


Parlatoria blanchardi, Targ. 
Aonidia blanchardi, Targioni Tozzetti, Mem. Soc. Zool. France, v, pp. 69-82, 
1892 (from Ourir, Shott Melrir, Algerian Sahara). 

Parlatoria victriz, Cockerell, Arizona Exp. Stn. Bull. 14, 1895 ; and Entomologist 
xxix, p. 52, 1896 (on imported date palms). 

? Parlatoria proteus var. palmae, Maskell, Trans. and Proc. New Zeal. Inst. (1897) 
Xxx, p. 229, 1898 (on date palms imported from Algeria into Australia). 

Parlatoria (Websteriella) blanchardi, (Targ.), H. H. King, Rept. Wellcome 
Res. Labs. Khartoum, iii, p. 240, 1908. 


296 P. A. BUXTON. 


Cockerell himself states that his description of Parlatoria victrix was made from 
dark specimens of the species previously named Aonidia blanchardi by Targioni 
Tozzetti. 


Distribution.—This pest has been recorded from the following localities : 


ALGERIA: Ourir, 8. of Shott Melrir. Eqaypr: Assouan, Cairo, Luxor. Supan: 
Khartoum and WN. Sudan generally. Iranian SoMALitaAnD: Merka. 
MesoporaMIA: Probably throughout the date area; certainly Basra and Amara. 
Inpia: Lyallpur, Punjab. Uwnirep States: Arizona, California. AUSTRALIA : 
Northern Territory. 


I can find no record of the occurrence of this scale-insect in Morocco, Tunisia, 
Tripoh, Arabia, or 8. Persia. 


P. blanchardi was introduced into the Southern States of the U.S.A. in 1890 
from Cairo and Algiers (Cockerell), or in 1889 (Popenoe). It was introduced into 
Australia about 1894 (Maskell), but has apparently not yet reached South Africa. 
(Lounsbury). 


Biology.—In Mesopotamia this scale occurs on the upper side of the leaflets, 
more rarely on the midrib; it attacks particularly recently planted off-sets and 
the resultant young trees during the first half dozen years of their life. I saw it 
on various types of date palms at and near Basra, and on the variety known as 
“ Zehedi’’ at Amara. I did not notice it in the other parts of Mesopotamia to 
which I travelled in the course of the investigation. I never saw a tree severely 
attacked, even in the most neglected gardens far from the river. I believe that the 
comparative scarcity of this insect is due to the attacks of Aphelinus mytilaspidis 
(EULOPHIDAE). 


Males were easily obtained in October by enclosing leaflets attacked by the 
scale in a bottle. 


This scale on the date leaf is known to the Mesopotamian peasant as “ remaj ”’ ; 
I have not been able to find this word in any dictionary. 


The most complete account of this scale is the pamphlet by Cockerell and Forbes 
(1907), to which I am indebted for a great deal of information. It is clear from 
all the American authors that it is a very serious pest, not only because it weakens 
the palm by almost covering the leaves, which turn yellow when heavily attacked, 
but also because it invades the fruit and renders the dates unsightly and hardly fit 
for the market. We know that it is also a very serious pest in the old world ; King 
speaks of dates exposed for sale in the Sudan “smothered with” P. blanchard, 
and Targioni Tozzetti’s original description of the species mentions date leaves 
from the Sahara covered on both surfaces by overlapping scales of this species. 
Essig says it may kill the tree in California. Cockerell (1907) states that it is 
attended by the ant, Phezdole cockerelli, Wheeler, at Tempe, Arizona. 


He and Forbes (1907) make the interesting suggestion that the young insects 
may be carried from tree to tree by wind, birds, or rats, or by fruit-pickers and 
pruners. Young offsets are frequently infested by both this and Phoenacococcus 
at the time when they are pushing their way through the old matted leaf-bases left 
on the trunk of the parent tree. 


INSECT PESTS OF DATES IN MESOPOTAMIA. 297 


Other Hosts.—Fide Cockerell (1907), Parlatoria blanchardi has been recorded 
by Newstead (1906) from jasmine foliage. King (1908) says Draper (1907) has 
recorded it from yellow jasmine and periwinkle. It does not appear to have been 
recorded from any other plant. 


Parasites and other Enemies —Cockerell (1907) records Coccinella abdominalis 
and Chilocorus cacti (COCCINELLIDAE) as feeding on this scale in America ; to this 
(Mthly. Bull. Sacramento, 1913, p. 139) we can add Scymnus sp. Popenoe states 
that there is a parasite which perforates this scale in Algeria, but it has not yet 
been investigated. No doubt this is some small Hymenopteron. Cockerell (1907) 
has once seen the emergence-hole of a Hymenopteron in a scale of P. blanchardi 
in America. 


In October 1918, at Amara, R. Tigris, in Mesopotamia, I bred a small Eulophid 
wasp from P. blanchard:. This parasite has been determined by Mr. J. Waterston 
as Aphelinus mytilaspidis, Baron. It was extremely common, and it appeared 
probable that it was an important agent in controlling the scale, which so far as 
I know is never a serious pest in Mesopotamia. It would probably not prove worth 
while to import this Eulophid into other countries in which the date is a main crop, 
because we know that the parasite is already widely distributed. According to 
Mercet, Aphelinus mytilaspidis has been found in U.S.A., Ceylon, Italy, France, 
and Spain, and is known to parasitize the following scales :—in America, Lepido- 
saphes ulma (Mytilaspis pomorum), Chionaspis pinifolir, and Diaspis carueli ; in 
Ceylon, Chionaspis permutans and C. graminis ; in Italy, Aspidiotus hederae and 
A. betulae; in Spain, A. hederae and Diaspis rosae. It has not apparently been 
previously recorded from Parlatoria blanchardi. 


Treatment.—Forbes (1907) states that treatment designed to exterminate Parla- 
torva blanchardi and Phoenicococcus marlatti has to be extremely thorough, because 
the scale-insects persist, even in small numbers, among the closely crowded bases 
of the great leaves. The reader must realize that the whole crown of the tree is 
enveloped in sheets of fibrous matting which arise between the petioles of the leaves 
where they are inserted on the trunk. Any scale-insects which happen to be in this 
tightly-bound tangle escape soap and kerosene emulsion, whale-oil soap, distillate 
spray, resin wash and even fumigation with hydrocyanic acid vapour. There re- 
main two treatments which are said to be efficacious. Wilsie (1913) tells us that 
after the great fire in San Francisco “the ornamental palms withstood the great 
heat, and put out new leaves at once after the fire.” This observation suggested 
the treatment described by Forbes (1913) ; the trees are pruned down to the crown 
and all the great leaf bases, and the fibre in which they are set, are removed as much 
as possible. When the tree has been reduced to a bare pole it is drenched with 
paraffin and fired, after which the surface is flamed with a gasolene blast, several 
times in the first few months, in order to destroy such few pests as may be finding 
refuge in cracks and similar places. This treatment does not kill the tree but des- 
troys the Parlatoria and Phoenicococcus. Popenoe (1913) says that the gasolene torch 
treatment puts the young palm back two years and is unpopular with the growers 
for that reason. He states that all offsets are now dipped bodily in a tank of cresol 
dip, and adds that “ this destroys the scale so that no danger need be feared from 


298 P. A. BUXTON. 


it.’ A plate is given showing the process in operation. I have no experience 
of this method but do not understand how any dip can penetrate the spaces between 
the leaf bases. No one but those who have endeavoured to separate date leaves 
from the trunk or even from a small offset can realize how firmly the petiole is 
attached to the plant, and what an ideal refuge there is among all the fibrous material 
which surrounds the bases of the petioles. 


Parlatoria zizyphi, Lucas. 

Various early authors refer to this scale as a pest of the date palm, and an un- 
signed editorial article in Insect Life (1891) definitely says that “ specimens proved 
on comparison to be identical with P. zazyphi Lucas.”” There appear to be no more 
recent records of P. zzyphi from date palms, and the above record dates from 
before the publication of the description of P. blanchardi, to which it possibly refers. 


Phoenicococcus marlatti, Ckll. 

Phoenicococcus marlattr, T.D.A. Cockerell, Proc. Acad. Nat. Sci. Philadelphia 

(1899), 1900, p. 262 (from imported Algerian date palms). 
Sphaerococcus drapert, R. Newstead, Quart. Jl. Inst. Comm. Res. in Tropics, 
1906 (from Egypt). 

Prof. Newstead tells me in a letter that his S. draperi is certainly identical with 
Cockerell’s P. marlattz. 

Distribution —Egypt and Algeria, both the Tel or Mediterranean coastal plain 
and also the Saharan Oases. As an imported species, California and Arizona. 

Biology.—I failed to find this species in Mesopotamia. Very little seems to be 
known of its habits. Cockerell (1899) says that the female is 1-1} mm. long, 
and wine-red in colour; she secretes wax but makes no true scale ; the embryos 
grow to a large size in the body of the female. Marlatt found it, according to the 
same author, occurring packed in great numbers “in little cavities about 10 or 12 
mm. long by 4 or 5 broad on the midribs of the leaves, communicating with the 
air by a narrow longitudinal slit,” on date palms imported into America from Algeria. 
The slight swellings containing the insects are very inconspicuous, and very likely 
to be overlooked even on a close examination. Later Cockerell himself found it 
on the sides of the sheathing bases of the leaves ; in this case it was not beneath 
the epidermis, and Forbes (1907) speaks of it as remaining close to the trunk among 
the great overlapping leaf-bases. Wilsie speaks of it as completely covering the 
stalk of the inflorescence near its base, where it is completely concealed by the leaf- 
bases. Probably Marlatt’s original description of the Phoenzcococcus living in cavities 
on the leaf-base was taken from specimens which had started growing on very 
young leaves ; as the leaf grew the tissues might have swollen round the colony 
of the scale-insect in such a manner as very nearly to cover it. Trabut’s most 
interesting note describes a date palm disease known to the natives of the oases 
of Zibans in the Sahara as ‘“‘ khamedj”’: this is caused by a heavy infestation of 
the stem of the inflorescence by P. marlatti. The disease becomes evident in the 
late spring soon a ter fertilisation ; the whole inflorescence dries up and no fruit 
develops. This calamity happens to 5 per cent. of the trees, nearly the whole of 


INSECT PESTS OF DATES IN MESOPOTAMIA. 299 


which are to some extent attacked. The Arabs treat the condition by free excision 
of leaves down to the bole, and occasionally by removal of the terminal bud itself. 
Forbes (1913) records that this pest occasionally attacks exposed roots of date palms. 


The male of Phoenicococcus marlatti is unknown. 
No parasites or natural enemies of this scale have been recorded. 


Icerya aegyptiaca, Douglas. 

This scale-insect is recorded by Newstead (1917) from a young date palm at 
Zanzibar. So far as I am aware this is an isolated record. This species has a long 
list of host plants, and in Ceylon at any rate attacks various kinds of palms, some 
of which are badly infested. 


Selenaspidus (Aspidiotus) articulatus, Morg. 

The West Indian red scale has been recorded by Morstatt from ‘‘ Phoenix palm,” 
from German East Africa. Iam not perfectly certain whether this refers to the date 
palm, or to some wild member of the genus. In any case it is an isolated record 
of a Coccid which has been found on a variety of plants, including olive, rubber, 
citrus, coffee, bananas and cacao. 


Aspidiotus ostreaformis. Curtis. 
Aspidiotus hederae, Vallot. 
Chrysomphalus aonidum, L. 
Chrysomphalus aurantii, Maskell. 
Coccus hesperidum, L. 
These five scales are all general feeders ; they are all recorded by Essig as occasional 
minor pests of the date palm in California. 

Mr. HE. E. Green has been good enough to furnish me with the following list of 
CoccipaE which have been recorded from the genus Phoenix :-— 

Icerya aegyptiaca, and purchase. 

Phoenicococcus marlatt. 

Pseudococcus nvpae. 

Lecanvum hesperidum, tessellatum, oleae, and expansum. 

Diaspis phoenicis, and borsduvali. 

Chionasprs dilatata. 

Aspidiolus ostreaefornus, dictyospermt, ficus, rapax, lataniae, hederae, cyanophylli, 

and articulatus. 

Parlatorva blanchardi, proteus var. palmae, zizyphi, and pergandei. 

Frorima pellucida. 

Ischnaspis filiforms. 

ACARINA. 
Family TETRANYCHIDAE. 

Oligonychus (Paratetranychus) ? simplex, Banks. 

Mr. Hirst has examined some of my specimens of a small “red spider” from 
date palms and believes that they are the Paratetranychus simplex described by 


300 F, A. BUXTON. 


Banks (Journ. Ent. Zool. Claremont, vi, 1914); it is apparently impossible to feel 
certain on this point. Mr. Hirst (1920) has re-described the species and figured 
the penis and the palp of the female. 

This mite is a most serious pest of growing dates in all parts of Mesopotamia 
which I visited. It was first noticed in early July, when immense numbers were found 
spinning webs all over the clusters of growing dates. These webs are of considerable 
extent and stretch from date to date over the whole cluster; dust becomes 
entangled in the web and the dates beneath never ripen properly. At the time when 
healthy dates are ripe the affected ones are discoloured and scaly, and as they 
are scarcely fit to eat and quite unsaleable they are generally used as food for cattle 
and horses. These diseased clusters are for some reason attractive to the wasp 
Polistes hebraeus, which may constantly be seen hovering over them and crawling 
on the surface of the webs. I was quite unable to find why the wasp acted im this 
manner, but I should imagine that its doing so must be a factor in the spread of 
the mites from tree to tree ; the wasp does not frequent the clusters of ripening 
dates unless they are attacked by the mite. The disease is found on many varieties 
of dates, especially the Khadhrawi, Dairi and Zehedi ; in 1918 1t was a most serious 
pest of the first of these, which is a widely grown date of superior quality and was 
not attacked by the Gelechud moth. I have seen clusters of dates on trees of all 
ages attacked by this mite, but it was always to be observed that its ravages did 
not cause serious damage except to trees far from water or growing in ill-kept 
unweeded gardens. It is not confined solely to such trees; mdeed I have seen 
it on a Khadhrawi which was growing on the very margin of a tidal creek at Basra. 
The disease is known to the Arabs as Toz, Trab, and Ajaj, all of which words mean 
solely dust ; it is stated to come from Allah, but more immediately from the dust 
of the roads. 

OTHER INsEcT PEsTS. 

,Mr. H. H. King informs me that in some years the migratory locust (Schistocerca 
peregrina, Oliv.) eats all the leaves and fruit on date palms in the Sudan, and 
Popenoe mentions that locusts are occasionally troublesome in California and 
Arizona, and in Algeria. He gives no further information, but we may presume 
that the adults of some species devour the dates on the tree. He makes some 
even more indefinite remarks about ants and termites, and also mentions bees 
and wasps. I have alluded above to the fact that Polistes hebraeus frequents 
clusters of dates infested by the mite Oligonychus simplex, Banks. The Oriental 
hornet (Vespa orientalis) never, I believe, attacks fruit on the trees except that 
which has been already pecked by birds, but it devours the fruit exposed for sale 
in the bazaars, and is so large and abundant that it must cause appreciable loss 
in all parts of Mesopotamia in which dates are grown. 


ANIMAL AND BirpD PEstTs. 

Popenoe tells us that a horde of rats at Tempe, Arizona, once did great damage 
to the date crop, and mentions that gophers and jack rabbits eat the offsets; he 
states that at Baghdad a “ squirrel-like animal” eats the fruit. The Mammalia 
of Mesopotamia are now well-known, but I am not able to guess which animal he 
was referring to; probably it is the mongoose (Mungos persicus), which is common 


INSECT PESTS OF DATES IN MESOPOTAMIA, 301 


ee 


in the date gardens and is known in Arabic as “ palm-tree rat’’; the mongoose 
is no doubt largely a vegetable feeder, but I do not think that it is in any way a 
serious pest of the date. 

In Mesopotamia birds undoubtedly eat a very large number of dates just before 
they are picked: Hypocolius ampelinus has been observed in Baghdad gorging 
itself and damaging many more dates than it could devour, and the bulbul 
(Pycnonotus leuco is mesopotamia) and house-sparrow (Passer domesticus) are also 
destructive. Mr. H. H. King informs me that the Spanish Sparrow (Passer 
hispaniolensis) pecks young dates in the Sudan. Milne adds parrots and monkeys 
to the list. 


Bibliography. 


Buckuier, W. (1882). Natural History of Ephestia passulella.—Ent. Mo. Mag., 
xix, pp. 104-106. 

Burkitt, L. H. (1913). The Coconut Beetles, Oryctes rhinoceros and Rhyn- 
chophorus ferrugineus.—Gardens Bull. Straits Settlements, i, pp. 176-188. 

Buxton, P. A. (1918). Report on the Failure of the Date Crop of Mesopotamia 
in 1918.—Bull. no. 6, Agric. Directorate Mesop. Exped. Force. (Provisional 
and inaccurate). 

CHITTENDEN, F. H. (1911). The Fig Moth.—U.8. Dept. Agric., Ent. Bull. no. 104. 

CocKERELL, T. D. A. (1899). Some Notes on Coccidae.—Proc. Acad. Nat. Sci., 
Philadelphia, 1899 (pub. 1900), pp. 259-275. 

CocKERELL, T. D. A. (1907). The Scale Insects of the Date Palm.—Bull. no, 
56, Univ. Arizona Agric. Exp. Stn., 22 pp., 5 plates, 5 figs. 

Coox, A. J. (1914). Monthly Bull. Cal. State Commis. Hortic. Sacramento, 
i, pp. 440-441, 2 figs. 

CopeLAND, EH. B. (1914). The Coconut.—xiv and 212 pp., 23 pl. Macmillan 
& Co. 

Draper, W. (1907).* Notes on the injurious Scale Insects and Mealy Bugs of 
Kgypt.—28 pp., 16 pl. Cairo. 

Durrant, J. H. (1915). Myelois phoenicis, sp. nov., bred ee Dates in Algeria 
and in England.—Ent. Mo. Mag., li, pp. 303-304. 

Essie, E. O. (1915). Injurious and Beneficial Insects of California.—Suppt. to 
Mthly. Bull. Calif. State Comm. Hortic., 2nd edition. | 

Forses, R. H. (1907). The Extermination of Date Palm Scales.—Bull. no. 
56, Univ. Arizona Agric. Exp. Stn. 

Forsrs, R. H. (1913). The Gasoline Torch Treatment of Date Palm Scales.— 
Jl. Econ. Ent., Concord, vi, pp. 415-416. 

Fracker, 8. B. Classification of Lepidopterous Larvae.—Illinois Biol. Mono- 
graphs, u, July 1915. 

GuosH, C. C. (1912). Life-history of Indian Insects. I1].—The Rhinoceros 
Beetle and the Red or Palm Weevil.—Mem. Dept. Agric. India, Ent. Ser. ii, no. 10. 
(713) H 


302 P. A. BUXTON. 


GouaH, L. H. (1913).* Entomological Notes.—Agric. JL, Egypt, ui, pp. 103-106. 

Gouenu, L. H. (1917). Notes on an Ephestia, an Insect mjurious to Stored 
Dates in Khargeh Oasis.—Bull. Soc. Entom. d’ Egypte, pp. 133-140, 1 pl. 

Haceporn, M. (1913). Borkenkafer (Ipidae) welche tropische Nutzpflanzen 
beschadigen.—Der Tropenpflanzer, xvu, p. 43, etc. 

Hampson, Sir G. F. (1917). A Classification of the Pyralidae, Subfamily Gal- 
lerianae.—Novitates Zoologicae, xxiv, p. 36. 

Hirst, 8. Revision of the English Species of Red Spider (Genera Tetranychus 
and Oligonychus).—Proc. Zool. Soc. Lond., 1920, p. 49-60. 

Kine, H. H. (1908). Report on Economic Entomology.—Rept. Wellcome Res. 
Labs., Khartoum, iii, pp. 201-248. 

Lounssury, C. P. (1918). Union Dept. Agric., Division of Entomology, Annual 
Report, 1917-1918, jp. 87-107. 

Mackenna, J. (1918). Report on the Progress of Agriculture in India for 
1916-1917. 

Maske, W. M. (1898). Further Coccid Notes with Description of New Species, 
and Discussion of Points of Interest.—Trans. and Proc. New Zeal. Inst., xxx. 

Minne, D. (1918). The Date Palm (Phoenix dactylifera) and its Cultivation 
in the Punjab.—Punjab Govt., Simla, 3rd edition. | 

Morstatt, H. (1913). Liste schadlicher Insekten.—Der Pflanzer, ix, pp. 288-296. 

NewstTEAD, R. (1906).* Quart. Jl. Inst. Commercial Res. in Tropics, April 
1906. 

NewstTeEaD, R. (1917). Observations on Scale Insects (Coccidae) V.—Bull. Ent. 
Res., vii, pp. 125-134. 

Popenog, P. B. (1913). Date-growing in the Old World and the New.—316 
pp., 40 pL, Publ. Altadena, California, West India Gardes. 

Romanorfr, N. M. (1901). Mémoires sur les Lépidoptéres, vii. 

Smytu, HE. G. (1911). Report on the Fig Moth in Smyrna.—U.8. Dept. Agric., 
Ent. Bull. no. 104. : 

STEBBING, EH. P. (1903).* Ind. Mus. Notes, v, pp. 126-127. 

TarGioni Tozzerti (1892). Aonidia blanchardi, nouvelle espéce de cochenille 
du Datier du Sahara.—Mém. Soc. Zool., France, v, pp. 69-82. 

Toumey, J. W. (1895).* Notes on Scale Insects in Arizona.—Ariz. Exp. Stn. 
Bull. .no. 14. 

Traput, L. (1912). Sur une Maladie du Datier, le Khamedj ou Pourriture du 
Regime.—C.R. Acad. des Sci., cliv, p. 303. 

Wixuoooks, F. C. (1914). The Date-stone Beetle-—Bull. Soc. Ent. d’Egypte, 
vi, 1918, pp. 37-39. 

Wituiams, C. B. (1918). Notes on some Trinidad Thrips of Economic 
Importance.—Bull. Dept. Agric. Trinidad and Tobago, xvii, pp. 143-146. 

Wiusiz, W. EH. (1913). The Date Palm Scales and their Control—Monthly 
Bull. State Comm. Hort. Sacramento, ii, pp. 538-539. 

(Unsigned editorial on Scales &c. in Sacramento).—Ibid., ti, p. 139-140. 


* Articles marked with an asterisk have not been examined by myself personally. 


INSECT PESTS OF DATES IN MESOPOTAMIA, 303 


Postscript. 

As this paper goes to press | have had an opportunity of examining the MS. 
of a “ Note on Dates and Date Palms in Iraq” by Mr. V. H. W. Dowson, which 
will shortly be published by the Directorate of Agriculture, Mesopotamia. I have 
also seen the “ Administration Report of Agricultural Directorate for the Year 1919” 
(Baghdad Govt. Press, 1920), which contains the report of C. R. Wimshurst, 
Entomologist. From these sources I gather that a “‘ minute larva has been found 
in the inflorescence of a date palm immediately after its exit from the spathe ” ; 
this is presumably the newly hatched larva of the unknown Gelechiid. Spraying 
with cold water at intervals of a few days is reported to be a valuable means of 
controlling the red spider (Oligonychus). Silvanus surinamensis, L., and Laemo- 
phloeus sp. (Coleoptera, Cucujidae) are added to our list of pests of stored dates. 


(713) H2 


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EXPLANATION OF PLATE VIII. 


A bunch of dates photographed in early August when growth of the fruit had 
practically ceased and ripening was just commencing : 

a, pedicel from which all the dates have fallen owing to the attacks of the larva 
of a moth (Fam. Gelechiidae) ; as will be seen, the perianth falls as well as the 
unripe date. 

b, dates destroyed by the larva, but which have not yet fallen; notice their 
small size and spherical shape ; they are quite hard and dry ; such dates are known 
in Arabic as “ hashaf.”’ 


c, healthy dates. 


BuLi. ENT. RESEARCH. VoL. XI. PART 3. PLaTE VIII. 


305 


LON THE GENUS PHLEBOTOMUS.—PART IV. 
By R. Newsteap, F.R.S., 
The School of Tropical Medicine, Liverpool. 


= The following notes may be regarded chiefly as materials towards the study of 
the geographical distribution of this group of the bloodsucking midges, which 
are of so much interest from a medical standpoint. For the opportunity of 
examining the very large series of specimens (570) herein recorded, | have to thank 
Dr. Guy A. K. Marshall, Director of the Imperial Bureau of Entomology, who 
sent the collections made by Drs. A. Ingram and J. W. Scott Macfie, in the Gold 
Coast ; and also that formed by Capt. J. Waterston, R.A.M.C., in Macedonia. 
I tender my thanks also to Capt. P. A. Buxton, R.A.M.C., for the collections made 
by him at Amara, Mesopotam a; to Capt. H. W. Leatham, R.A.M.C., for examples 
from Basra; and to Major J. A. Sinton, V.C., I.M.S., for the specimens which he 
collected in N.E. Persia, under exceptionally difficult circumstances, and also 
for the long series taken by him in the N.W. Frontier Province of India. 

I may add to this short communication that the determinations of the specimens 
herein recorded are based upon material which was carefully prepared and stained 
for microscopical examination, as by this means the minute taxonomic characters 
can be seen and studied with greater accuracy and ease. 


Phiebotomus antennatus, Newstead. 

Male (hitherto unknown). Pale form. Drab or straw-coloured ; fringe of costa, 
anteriorly, rather strongly infuscated, that of the posterior margin not so, Hairs 
NS 


Fig. 1. Phlebotomus antennatus, Newst., g; a, proximal segments of the 

antenna; a1, terminalsegments. P. minutus africanus, Newst., $: b, proxi- 

mal segments of antenna. P. simillimus, Newst., J: ¢. proximal segments 

of antenna. The figures a, b, and ¢c are drawn to the same scale; ai, ata 
much higher magnification. 


on wings and scales on legs very faintly iridescent in a strong light. Dark form. 
Wing very heavily infuscated, especially the costal fringe on both margins ; the 
hairs beautifully iridescent, and in certain lights strongly resembling the rich 
deep blue of lapis-lazuli. The scales of the legs are also iridescent in certain lights, 


306 R. NEWSTEAD. 


the predominant blue colour being almost as intense as that of the wings. Hairs 
on the proximal segment of the abdomen erect ; those on the remaining segments 
recumbent, Antennae with the segments (fig. 1, a) only very slightly longer 
than those of the female ; the geniculated spines unilateral* and unpaired. Wing 
rather narrowly lanceolate. External armature precisely similar to that of 
P. minutus, Rond. 

Length, 2-2°4 mm.; wing 1'°2 mm.; leg 11, 2 mm. 

Gotp Coast: Accra, in laboratory. vi. 19, 38 §g,59 292 (Dr. J. W. Scott Macfie) ; 
Kwaman, 10.vii.18, 12; Navarro, on walls of M.O’s. quarters, 22.vi.18, 1 9; 
Yamalaga, 18.v.18, 1 2; Salaga, on walls of M.O’s. quarters, 10.v.18, 1 9; Yeji, 
6.v.18, 1 2 (Dr. A. Ingram). 


Phiebotomus ingrami, Newstead. 
Gotp Coast: Chechewere, 9.vii.18, 1 g (Dr. A. Ingram). 


Phlebotomus minutus, Rondani. 

Maceponta : near Salonica, 1918, 1g, 19 (Capt. J. Waterston). MESOPOTAMIA : 
Amara, River Tigris, 1.1x.18, 1 3, 20 9° (at light); 81x.18,2 gg, 5 2 (at 
light) 3" 19.1%-18, 1°95" 30.1%.18, 1 gy 9 YO lat heht): Tis eo: ea 
1 ¢ (at light) (Capt. P. A. Buaton). Inpt1a: N.W. Frontier Province, Dera 
Ismail Khan, vii. 1919, 33 gg, 34 92 (Major J. A. Sinton, V.C.). 


Phlebotomus minutus africanus, Newst. 

Gotp Coast: Yeji, 6.v.18, 1 9; Malowe, 29.vi.18, 1 g, 299; Accra, 4.1.19, 
2 ¢3, 1 9; Lilxia, 4.vi.18, 3 gg, 3 29; Diari, on Rest House wall, 26.v.18, 
1 2; Gambaga, in latrine, 1.vi.18, 7 gg, 1 9; 5.vi.18, 1 2 inside mosquito 
net ; Salaga, on walls of M.O’s. quarters, 10.v.18, 6 gg, 3 Q2; Jugbe, 1.viu. 18, 
1 2; Sekodumase, 8.vii.18, 2 gg, 11 292; Til, on walls of Rest House, 
14.vi.18, 2 §g, 12; Nangudi, on walls of Rest House, 15.vi.18, 3 dg, 7 99; Kul- 
masa, 22.vii.18, 1 9; Nandawli, 15.vu.18, 2 gg, 429; Bole, 26.vi.18, 1 ¢g; 
Chechewere, 9.vii.18, 1 g, 3 29; Bawku, 12.vi.18, mside mosquito net, 3 99; 
Savelugu, Rest House, 25.v.18, 2 3g, 2 99; Wa, Rest House, 20.vu.18, 1 9; 
Navarro, on walls of M.O’s. quarters, 22.vi.18, 2 3g, 4 99; Bawku, on walls 
of quarters formerly belonging to M.O., 10.vi.18, 5 gg, 1 2; Banda Nkwanta, 
30.vu.18, 2 gg, 6 29; Maibindiga, on walls of Rest House, 3l.v.18, 2 99; 
Palbe, 16.v.18, 1 9; Kwaman, 10.vili.18, 1 9; Zovaragu, on walls of D.C’s. 
quarters, 16.vu.18, 1 g, 2 99; Tishi, on walls of Rest House, 1.vi.18, 1 g, 1 9; 
Kpalgu, on walls of Rest House, 30.v.18, 1 9; Yamalaga, 18.v.18, 1 2 (Dr. 
A. Ingram); Accra, in laboratory, vi.19, 3 3g, 4 992 (Dr. J. W. Scott Macfie). 


Phiebotomus papatasii (Scopoli). 

MrsopoTaMIA: 30.vi.18, 1 3, 3 99; Amara, River Tigris, 14.v1.18, 1 9; 
vili,18, 1 6, 5 O93 2.1%:18,3 99 (biting man); 211x18, 1. og (Cap F. A. 
Buxton). Macrponta: near Salonica, 1918, 14 gg, 35 29 (Capt. J. Watersion). 
Inp1a : N.W. Frontier Province, Dera Ismail Khan, vii. 1919, 3 ¢¢, 3 22 (Major 
J. A. Sinton, V.C.). Asta Minor: Trebizond, 5 gg, 3 99 (Dr. B. Uvarov). 

* This rather remarkable character is common also to the males of the following 


species: P. minutus, Rond., P. minutus africanus, Newst., and P. simillimus, Newst. 
(see fig. 1 b, ¢). 


ON THE GENUS PHLEBOTOMUS.—PART IV. 307 


Phlebotomus perniciosus, Newstead. 


MaceponliA: near Salonica, 1918, 26 3g, 9 2° (Capt. J. Waterston), CaucasiA: 
Borzham, Province of Tiflis, 16.vii.1919, 8 gg, 3 22 (Dr. Kandelaki, per 
Dr. B. Uvarov). 


Phiebotomus sergenti, Parrot. 


Phlebotomus sergenti, Parrot, Bull. Soc. Path. Exot. x, pp. 564-567, figs. 1-11 
(1917). 


Phlebotomus caucasicus, Marzinovsky, Medical Review, Moscow, lxxxvii, nos, 
13-14-15-16, pp. 612-614 (1917). 

A large species somewhat resembling P. papatasi (Scop.) in colour and in the 
general arrangement of the body hairs. But the male of P. sergenti is abundantly 
distinct and may be recognised from all other species by the unique character 
of the genitalia, in which the great length of the sub-proximal spine on the superior 
claspers should serve at once to distinguish it from all the species possessing paired 
brushes (peniculz) of non-deciduous hairs between the superior claspers proximally. 
On the other hand, it is extremely difficult to give taxonomic characters for the 
separation of the female from that of P. papatasw. Ina typical example of P. sergentt, 
generously presented to me by M. Parrot, I fail to find any structural difference 
by which it could be separated from the 9 of P. papatasw, though it is decidedly 
smaller ; and this applies also to the female of the variety herein described from 
Mesopotamia. In the females from Persia, however, the geniculated spines of 
the antennae are slightly longer than those on the corresponding segments in the 
African examples before me ; and in one example the spines are also accompanied 
by a somewhat lunular-shaped, subcutaneous structure (? gland), which no staining 
has hitherto revealed in any of my preparations. On the other hand, the males 
from the two widely separated regions agree in all their morphological details 
and are clearly conspecific. There is, I think, no doubt also that the African and 
Caucasian species are one and the same. 

Marzinovsky (l.c.) notes the close affinity of his P. caucasicus with P. sergentz, 
Parrot, but claims that it is distinguished from the latter in some details. These 
‘* details ’ are, however, not traceable either in his diagnosis or in his figure of the 
male genital armature. Thanks to Col. C. M. Wenyon I have been able to study 
a direct photographic print of the male genital armature taken from the original 
negative which was used by Marzinovsky in illustration of his paper, and this con- 
firms my views as to the specific identity of the Caucasian with the African form. 

The following description is based upon material collected in N.E. Persia by Major 
J. A. Sinton, V.C. :— 

Male.—Palpi of five segments; 2nd and 4th about equal in length; the 3rd 
a little longer than the 2nd, laterally incrassate, the incrassation seen only in dorso- 
ventral view ; sensorium well marked by the relatively large clubbed hairs ; 5th 
a little more than twice the length of the 4th. Antennae with the distal end of 
the 3rd segment almost reaching to the tip of the proboscis ; the tips of the paired 
geniculated spines on segments 3-15, inclusive, reaching a point very slightly 
in advance of the centre of all the segments with the exception of the 3rd. Wing 


308 R. NEWSTEAD. 


(fig. 2, a) somewhat lanceolate, the tip rather acutely pointed. Genital armature 
(fig. 2, 6) with a pair of relatively large brushes or peniculi between the superior 
claspers proximally. Proximal segment of superior claspers very robust; distal 
segment about half the length of the first, strongly bifurcated distally, the tip of each 


Jmm Vega iar eee Oa 5 ates 


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Fig. 2. Phlebotomus sergenti, Parrot: a, wing of J; b, genital armature of ¢ ; 
c, sixth segment of antenna of 9, showing relative length of the geniculated spines 
and subcutaneous structures (? glands) near the bases of the former; ci, dotted 
outline of the corresponding segment in P. papatasii ; d, wing of P. sergenti 2 , in 
dotted outline, drawn to same scale as that of thed (a); e, connective cuticle 


_ between tergites and sternites. 


branch with an unusually long stout spine ; near the proximal third of the segment 
is another pair of spines: the outer lateral one stout and of great length, the tip 
of which reaches almost to the base of the proximal segment of the superior clasper 
when the latter is closed (see fig. 2, b) ; the inner lateral spine much more slender, 


ON THE GENUS PHLEBOTOMUS.—PART IV. 309 


generally curved distally and a little more than half the length of the longer outer 
one. The long inferior clasper extending beyond the distal end of the proximal 
segment of the superior clasper by one-fifth of its total length. 

_ Length, 3 mm ; length of wing, 1°9 mm. 

Female.—General facies similar to that of the male, but larger and generally 
more robust. Palpi precisely similar to those of the male. Antennae with the 
tip of the 4th segment extending very slightly beyond the proboscis; the 3rd 
segment relatively shorter than the corresponding segment in the male ; the paired 
geniculated spines (fig. 2, c) extending considerably beyond the middle in most 
of the segments, while those on the 14th and 15th, respectively, extend to the 
articulation of the succeeding segment ; on many of the segments the spines are 
accompanied by a relatively large and somewhat lunular-shaped subcutaneous 
structure (? gland). Wing (fig. 2, d) much broader than that of the male, with the 
hind margin also much more strongly arched. Connective cuticle between the 
tergites and sternites (fig. 2, e) with a very regular diagonal pattern formed by 
equidistant groups of minute hairs, each group composed of four hairs. 

Length, 3°7 mm. ; length of wing, 2°4 mm. ; length of leg u, 3°9 mm. ; length of 
3rd segment of antennae, 0°3 mm. 
ae Kast Persia: Meshed, vi. 1918, 5 gg, 3 99 (Major J. A. Sinton, V.C.). 

as 
Phiebotomus sergenti, var. 

Male.—Difiering from typical P. sergenti, Parrot (l.c.) in having the 3rd segment 

of the antennae (fig. 3, a) relatively very much shorter and with the distal, bifur- 
icating, spine-bearing process of the superior claspers (fig., 3 6) markedly unequal 
in length, the terminal process being about three times the length of the other. 


Fig. 3. Phlebotomus sergenti, var. 3: a, proximal segment of the antennae ; 
b, b, distal segments of the superior claspers. 
Phlebotomus sergenti, Parrot, 3: ¢, proximal segments of the antenna; d, 
distal segment of the superior clasper, 
All the figures are drawn to the same scale. 


310 R. NEWSTEAD. 


The 3rd antennal segment (fig. 3, c) and the distal segment of the superior clasper 
(fig. 3, d) of P. sergents drawn to the same scale as the corresponding structures 
in this var. are given for comparison and convenience of reference. 


Mesopotamia: Amara, 1 dg, taken at light, 19.1x.18 (Capt. P. A. Buaton) ; 
another ¢ from the same locality, but taken 1.ix.18, has been placed in the National 
Collection. There are also 6 99 which I attribute to this variety of P. sergenit, 
though I am not absolutely certain that they belong here ; they bear the following 
data: Amara, River Tigris, 4.vi.18, 1 9; Mesopotamia, 30.vi.18, 3 99; Baqubah, 
River Diayala, 28.vii.18, 1 9; Amara, River Tigris, 18.x.18, 1 9 (biting man) 
(Capt. P. A. Bucton). 


Phiebotomus signatipennis, sp. nov. 

A small short-legged exceptionally hairy species; easily determined by the 
strikingly characteristic wing venation, in which the Ist sub-marginal cell is 
relatively very small indeed, the anterior branch of the vein being about one-fifth 
the length of the vein between the two forks. 


Fig. 4. Phlebotomus signatipennis, Newstead, 2: a, palpus; 
b, wing ; the latter drawn to the same scale as the wing of P. sergenti 
in fig. 2, a. 


Female. Relatively robust, short-legged, and exceptionally hairy, the hairs 
heavily infuscated, more especially those which form the wing-fringe. Hairs on 
proximal portion of abdomen more or less erect, the longest reaching to the middle 
of the 4th segment, the rest recumbent on both the tergites and sternites. Leg 
scales dark leaden grey ; hind tibiae with 4-5 long equidistant outstanding hairs. 
Antennae with relatively short segments, and densely clothed with long hairs. 
Palpi (fig. 4, a) relatively short and robust; 2nd segment rather densely clothed 
with very long hairs, the longest reaching to about the middle of the succeeding 
segment; 3rd strongly incrassate and about twice the length of the 2nd; 4th 
narrower, and a little shorter than the 3rd; 5th relatively very short and about 
equal in length to the 3rd. Clypeus very short, and widely rounded distally. 
W_ngs (fig. 4, 6) narrowly lanceolate, and distinctly pointed; veins unusually 
thick; the 1st sub-marginal cell exceptionally small, and placed at a relatively 
long distance in advance of the tip of the 1st longitudinal vein, the distance 


ON THE GENUS PHLEBOTOMUS.—PART IV. 311 


between the two forks of the 2nd longitudinal vein being five times as great as 
the length of the upper branch of the 2nd fork, hairs forming the fringe dense and 
of great length, some of them being much longer than the greatest width of the 
wing. The hairs and scales on all parts of the body are not, apparently, so easily 
deciduous as in other members of the genus. 

Length, 16 mm, approximately; length of wing, 1:25 mm. 

Goitp Coast: Gambaga, in latrine, 1 9 (type), 1.vi.18 (Dr. A. Ingram). 


Phiebotomus simillimus, Newstead. 

Gop Coast: Yamalaga, 18.v.18, 1 g,3 92; Kpalgu, 30.v.18, 1 g¢; Kukuma: 
6.viiil.18, 1 g, 1 2; Maibindiga, on walls of Rest House, 31.v.18, 1 ¢; Nkoranza, 
9.viii18, 2 gg, 10 99; Kulmasa, 22.vi.18, 1 9; Sekodumase, 5 99 Yeji, 
8.viil.18,; 6.v.18, 1 g (Dr. A. Ingram). 


Phiebotomus walkeri, Newstead. 

Phlebotomus walkeri, Newst., Bull. Ent. Res. v, p. 190 (1914). 

Phlebotomus longipalpis, Newst. (nec Lutz & Neiva), abid. p. 188. 

The species taken by Dr. F. D. Walker on the Abuna River, Bolivia-Brazil 
Boundary, in 1913, and recorded by me in this Bulletin (l.c.) as P. longipalyis, Lutz 
& Neiva, is, as I surmised at the time, clearly distinct and, so far as I can judge, 
must take specific rank under the alternative name, walkeri, suggested by me 
(ie)< 

The males of the two species are easily separable by the characters of the external 
genitalia: in P. longvpalpis the small brushes or peniculi are absent, and the 
four long spines on the distal segment of the superior claspers are arranged as 
follows: one apical, one sub-apical, and the remaining two placed closely together 
on opposite sides of the segment in a similar way to the corresponding spines in 
P. sergenti (see fig. 2, 6) ; whereas in P. walkeri the 4th spine is quite isolated from 
the others. 

I am pleased to be able to clear up the discrepancies to which I had previously 
called attention, thanks to the generosity of Dr. Lutz, who has very kindly presented 
me with examples of his P. longipalpis. 


Phlebotomus sp. 

The examples recorded below come very near P. antennatus, Newst., and possibly 
represent a degree of variation in the antennal formula of the females of this 
species. But pairs taken in coitu are needed before one can state to what extent 
variation occurs in the females of this and other allied species. 

Gotp Coast: Yeji, 6.v.18, 1 9; Savelugu, in Rest House, 25.v.18, 1 2; Wa, 
on walls of Rest House, 20.vu.18, 1 9; Navarro, on walls of M.O.’s quarters, 
22.v1.18, 2 99; Kwaman, 10.vii.18, 3 99; Yamalaga, 18.v.18, 2 99; Salaga, 
on walls of M.O.’s quarters, 10.v.18, 1 Q (Dr. A. Ingram). 


Phiebotomus sp. 
Mesopotamia: Amara, x.18, 1 Q (at light); 81ix.18, 1 9 (at light); Amara, 
River Tigris, 30.ix.18, 1 9 (Capt. P. A. Buaton). 


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313 


COLLECTIONS RECEIVED. 


The following collections were received by the Imperial Bureau of Entomology 
between Ist July and 30th September, 1920, and the thanks of the Managing 
Committee are tendered to the contributors for their kind assistance :— 

Mr. E. Ballard, Government Entomologist, Madras :—1 Tachinid fly, 17 Coleop- 
tera, and 28 Lepidoptera ; from South India. 

Captain P. J. Barraud, Entomologist to the Egyptian Expeditionary Force :— 
235 Culicidae, 1 Phlebotomus, 7 Tabanidae, 1 Hippoboscid, 84 other Diptera, 39 
Hymenoptera, 69 Coleoptera, 2 Planipennia, 13 Lepidoptera, 5 Isoptera, 18 
Rhynchota, 6 Orthoptera, 34 Odonata, and 1 Tick ; from Palestine. 

Mr. John R. Bovell, Superintendent of Agriculture :—3 Diptera, 22 Hymenoptera, 
2 Coleoptera, 6 Lepidoptera, 4 Thysanoptera, 9 Isoptera, and 15 Mites; from 
Barbados. 

Dr. Briinn, Jewish Health Bureau :—A series of insects affecting Cuscuta ; 
from Palestine. 

Mr. P. A. Buxton :—315 Orthoptera; from Mesopotamia. 

Mr. P. R. Dupont, Curator of the Botanic Station :—3 Diptera, 21 Ants, 1 Moth, 
1 species of Coccidae, and 100 Mites; from Seychelles. 

Mr. T. B. Fletcher, Imperial Entomologist, India :—134 packages of Coccidae ; 
from India. 

Mr. G. F. Hill, Entomologist, Australian Institute of Tropical Medicine :— 
17 Culicidae, 8 Tabanus, 4 other Diptera, 3 species of Coccidae, and 3 other 
Rhynchota; from North Queensland. 

Mr. M. Afzal Hussain, Government Entomologist, Punjab :—21 Diptera ; from 
the Punjab. 

Mr. J. C. Hutson, Government Entomologist :—28 Coleoptera, 47 Lepidoptera, 
25 Rhynchota, and 470 Ticks; from Ceylon. 

Dr. J. F. Ilingworth :—188 Hymenoptera ; from Queensland. 

Dr. W. A. Lamborn :—41 Culicidae, 17 other Diptera, 3 Moths, 27 Lepidopterous 
puparia, and 236 Chalcids ; from Nyasaland. 

Dr. Ll Lloyd :—3 tubes of Aleurodidae ; from Herts. 

Dr. R. EH. McConnell :—13 Tabanidae, 244 Glossina, 3 Stomoxys, 13 other 
Diptera, 12 Hymenoptera, 27 Coleoptera, 14 Rhynchota, and 7 Orthoptera ; from 
Uganda. 

Prof. G. H. F. Nuttall, F.R.S. :—7 Tabanus, 62 Hippoboscidae, and 100 early 
stages, 20 other Diptera, 54 Coleoptera, 2 Hymenoptera, 4 Rhynchota, and 2 
Anoplura ; from various localities, 

The Rev. Father J. A. O’Neil, S.J. :—75 Coleoptera and 13 Moths; from 
Rhodesia. 

Mr, A. W. J. Pomeroy, Government Entomologist :—6 Tabanus, 34 other 
Diptera, 25 Chalcids, 40 other Hymenoptera, 78 Coleoptera, 34 Lepidoptera, 36 
Rhynchota, and 4 Planipennia ; from Southern Nigeria. 


314 COLLECTIONS RECEIVED. 


Mr. A. H. Ritchie :—1 species of Coccidae ; from Jamaica. 

Senhor A. F. de Seabra :—37 Coleoptera ; from San Thomé. 

Mr. Hubert W. Simmonds :—5 Diptera, 3 Lepidoptera, 1 Lace-wing, and 8 
Chalcids ; from Polynesia. 

Mr. S. H. Skaife, Government Entomologist :—2 Diptera and 34 Coleoptera ; 
from Natal. 

Mr. R. Swainson-Hall :—2 Beetle larvae ; from Portuguese East Africa. 

Mr. O. H. Swezey, Entomologist to the Hawaiian Sugar Planters’ Association :— 
9 Coleoptera, and 11 Rhynchota; from Hawai. 

Mr. Robert Veitch :—54 Diptera, 140 Chalcids, 16 other Hymenoptera, 1,440 
Coleoptera, 85 Lepidoptera, 1 Caddis-fly, 72 Rhynchota, 4 Orthoptera, and 1 
Chrysopa: from Fiji. 

Wellcome Bureau of Scientific Research :—15 Tabanidae, 22 other Diptera, 
2 Hymenoptera, and 7 Rhynchota; from various localities. 

Mr. R. O. Williams, Superintendent of Agriculture, Grenada :—Material of 
leaf blister mites ; from Carriacou and Isle d’Rhonde. 


— IMPER IAL ‘BUREAD OF ENTOMOLOGY. 
HONORARY COMMITTEE oF MANAGEMENT 


“THE VISCOUNT HARCOURT, Cbairman. |” Bye 
 Lreur.-Cononetr A. ALCOCK, C.LE., F.R.S. ea 
Mayor H. E. AUSTEN, D.S.0. - | 
Dr. A. G. BAGSHAWE, OMG. i 
Masor-GeneRAL Stn JOHN R.-BRADFORD, K. CMG. ERS. 
Masgor-Generat Sir ‘DAVID BRUCE, K.CB., PRS. | 
Mr. J. C. F. FRYER. | 
‘Sir & F. HARMER, K.B.E., ERS. " 
. Pror. H. MAXWELL LEFROY. 
Hon. E. LUCAS. § 
Dr. R. STEWART MACDOUGALL. 
Sir JOHN McFADYEAN. 
Sin PATRICK MANSON, G.O.MG., F.R.B. 
‘Sim DANIEL MORRIS, K.0.M.G. 
- Pror. R. NEWSTEAD, F-.R.S.. GE Ne ea Ch ence OME 
Pror. G. H. F. NUTTALL, F.R.S. oa apa name tat | 
Pror. E. B. POULTON, E.R. . V6 
Lizvut.-CoLonEL Sie DAVID PRAIN, O.M.G., OLE, PRS. 
Sm H. J. READ, K.C.M.G,, OB. ' 
Tun Hox. N. @. ROTHSCHILD. = = 9° 
Dr. HUGH SCOTT. Be 
Sin A. B. SHIPLEY, GBE,, PRS. 
‘Me. R. A.C. SPERLING, OMG. © . Bah 
Sm STEWART STOCKMAN. TA ane 
Me. F. V. THEOBALD. 3 | Beakicka | 
Mr. C. WARBURTON. | 


I 
ch 


i | Director. Ah i Mite 
| Dr Guy A. K, MARSHALL, ome, Nice 


_Bosistant. Director. 
‘SB. Ae NEAVE, — 


:  Gecretaty. ae Comey 
Mr, A. 0. ©. PARKINSON. ie. 


{ ¢ 
My a ie 
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315 


AN EXAMINATION OF THE TSETSE PROBLEM IN NORTH MOSSURISE, 


PORTUGUESE EAST AFRICA. 
By C. F. M. Swynnerton, F.LS. 
(Puates [X—XVII and Map.) 


CONTENTS. 

PAGE. 
T. Introduction .. 315 
Il. Nature of the Country ir cichdepatia :«/ S48 
III. Wooding of the District analysed in sheticn to T sateen pong 
IV. Factors influencing Leaf-fall Fee des’ .. d20 
V. Effect of the annual Grass-fires .. 324 
VI. Distribution of Rocks, Woodland Types, Food Actnsial sad Hy 1 325 
VIL. Umzila’s Experiments in Close Settlement be .. 331 
VIII. The Food of the Fly . -» 3086 
IX. Distributors of the Fly . 342 
X. Effect of the Rinderpest 9343 
XI. Factors determining the Presence of Dashed . 344 
XII. Notes on Habits .. 355 
XIII. Proportion of the Sexes , 36] 
XIV. Observations on Breeding Habits sm» OO 
XV. Net Experiments (a BBS 
XVI. Natural Enemies . 0908 
XVII. Tsetse Concentrations by 369 
XVIII. The Recent Outbreaks of Nagana in n Mossurise sai ee 
XIX. Clearing Experiments m sia 
XX. Measures for the Control of igetees 1 in Mossuriae . 375 
XX]. Discussion of General Methods of Tsetse Control . a» G40 


I.— INTRODUCTION. 


In June 1918, I arranged to carry out a preliminary investigation of the habits 
and distribution of the tsetse-flies in the northern portion of the Mossurise district 
for a maximum period of three months on the understanding that the Mozambique 
Company would defray the general expenses of the expedition. 

After a brief stay at Gogoyo’s with little success I proceeded to the country east 


of the Sitatonga Hills. 


I had found tsetses in considerable numbers there in June 


1900, and hoped by commencing my work in a well infested area to gain at once 
a thorough acquaintance with the fly that would stand me in good stead during 
the rest of my expedition and enable me also to decide quickly on my requirements 
in the matter of bait and labour. I was fortunate enough to make a number of 
particularly interesting observations during this first part of my trip, east of the 
Sitatongas. 

Having found that the large tsetse (Glossina brevipalpis) does not come readily 
to man or to goats, | applied for cattle to serve as bait. These reached me on 
(737) 1,000. Wt.P2/154. 3.21. B.&F.,Ltd. Gp. 11. A 


~—-316 C. F. M. SWYNNERTON. 


28th July, and proved quite invaluable during the remaining six weeks. They 
revealed abundant fly in places in which the nature of the vegetation had previously 
been the only indication of its possible presence, and led to the suggestion that one 
reason why investigators have commonly met with so enormous a preponderance 
of male flies is that insufficiently attractive bait has been offered to the sex that 
comes only to feed. 

The part of my investigation already referred to, east of the Sitatonga Hills, 
was carried out with the help of my own farm natives only, and was concerned 
chiefly with the small flies, Glossina morsitans and G. pallidipes, which I there found 
mixed. I also then made my discovery, and commenced my study, of the Mtsha- 
nedzi-Buzi brevipalpis area, which at its other end affects Spungabera. 

The second month (I was now being assisted in the matter of labour by Snr. 
Lanne) was occupied mainly with a fly survey of the northern part of thé district. 
It included also the completion of my work on the breeding habits of G. morsitans 
and a week’s study, south-west of Chibabava, of a typical piece of the surface-basalt 
country that constitutes so very large a proportion of the low-lying parts of the 
Mossurise district. At this period I sent a successful side-expedition round north 
of the entire Lusitu River in the Moribane district in order to gain some idea of the 
full extent and importance of the eastern fly-areas. 

The third section of the trip was occupied with the completion of the above work, 
with experiments directed to ascertaining the effect and cost of particular forms of 
clearing for fly, a study of the habits of G. brevipalpis and a special study of the 
Spungabera problem. Spungabera being the official head-quarters of the district, 
and the seat of a herd of cattle belonging to the Mozambique Company that had 
lost heavily from trypanosomiasis, its protection from fly appeared to be of the 
first importance. 

{ returned home finally on 10th September, and afterwards I sent a small 
expedition to the Chinyika River, south of Spungabera, and completed my 
breeding experiments. 

Four days had been spent near the British border examining the details of nagana 
outbreaks in Portuguese territory, two in trying to ascertain the maximum 
dry season elevation of G. brevipalpis, and two in all at Spungabera itself. Otherwise 
all the work was more or less far within Portuguese territory. I had had four 
working camps—one on the Umvuazi River east of the Sitatongas, one on the 
Buzi in the Gunye-Umpombo country south-west of Chibabava, one near the east 
end of the great southern brevipalpis area, and-one at its west end at a point on the 
Buzi six miles from Spungabera. 

The Mossurise fly problem quickly proved to be of quite extraordinary interest. 
Apart from the presence in the district of so many different tsetses, and the oppor- 
tunity of studying them usefully that is offered by the very varied but well- 
demarcated conditions of vegetation, elevation, rock-formation, etc., that the district 
afiords, the area west of the Sitatonga Hills was, under the Zulu domination, the 
scene of a particularly fine experiment in the banishment of tsetse. It was not 
difficult to obtain the details of this experiment, as most of the older natives had, 
incidentally or by compulsion, taken part in it. I made particular efforts to 
obtain information on this point, (a) because a study of any successful campaign, 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 317 


by whomsoever carried out, is bound to add usefully to our general knowledge on 
the subject of controlling tsetse ; (b) because the area cleared of fly was the very area 
which, in the Mossurise district, is best adapted to white settlement. This experi- 
ment affords a very clear indication that settlement, properly planned, is itself 
capable of clearing the country settled of the two flies here concerned—G. brevipalpis 
and G. pallidupes. 

Other points to which I paid particular attention were an analysis of the vege- 
tation in relation to tsetse, and a study of the distribution and local behaviour 
of the dangerous and less dangerous woodland types. The practical study of tsetse 
is, 1 am convinced, a matter for the botanist and cecologist rather than the unaided 
entomologist, and this point will be seen to be of the first importance. Hach of 
the tsetses is dependent on particular, but different, shade conditions. Without 
a clear knowledge (a) of the woodland types that provide these conditions, (b) of 
those that provide them throughout the year, and (c) of the dry season response 
of each of these types to elevational and edaphic conditions, exceptional seasons, 
burning, etc., any clearing operations that may be undertaken in the future for the 
extirpation of fly, in connection with settlement or otherwise, will be lacking 
completely in the elements of precision and economy. 

The main practical results have been, I think, the clear defining of the several 
fly-areas and of the situation at Spungabera, the lessons, both general and in relation 
to future settlement in Mossurise, to be drawn from the details of Umzila’s success, 
and, in relation to G. brevipalpis in particular, the fact that the annual grass-fires 
can be made to play an important part by regulation of the time of burning. In 
primary forest the control of this fly is unfortunately rendered difficult by the 
presence of the Landolphia vines, which it may not be wished to sacrifice. 

So far as G. morsitans is concerned, it seems to me that the possibility of taking 
advantage of its highly gregarious and localised habits is worthy of further study. 

I conclude by expressing my very high indebtedness to Snr. Lanne and, in his 
absence, Snr. A. P. Monteiro for going out of their way to do all in their power to 
assist me in making a success of the investigation, and to my old friend Snr. Ferreira 
for his very cordial offer of help on learning that I was working in his district also. 
I am likewise particularly highly indebted to Dr. W. T. Lawrence of Gogoyo, who 
not only gave me such information as he already possessed with regard to the fly 
and continuously collected tsetses for me locally, but also rendered my work east 
of the Sitatongas and south of the Mtshanedzi possible in this foodless season by 
lending me native food in considerable quantity. Mrs. Lawrence most kindly 
presented me with gauze for nets at a moment when it was particularly acceptable, 
and Messrs. J. W. Scott and G. D. Otterson, living in the mountains, were good 
enough to assist me in a test of the resistance of G. morsitans to cold in the pupal 
stage. For some of the literature of the subject, presented to me at different periods 
in the expressed hope that I would undertake tsetse investigation, I am indebted to 
my friends Drs. G. A. K. Marshall and A. G. Bagshawe, Directors of the Imperial 
Bureau of Entomology and the Tropical Diseases Bureau, London, respectively ; 
and for copies of his own interesting publications, to Mr. R. W. Jack, Rhodesian 
Government Entomologist. I have also to thank the Rhodesia Museum and Mr 
F. P. Mennell, of Bulawayo, for the latter’s identifications of a rock collection 

(737) A2 


318 C. F. M. SWYNNERTON. 


made in connection with the tsetse work ; Mr. E. C. Chubb, Curator of the Durban 
Museum, for his identification, still incomplete, of the small mammals trapped 
by me in a dry season breeding centre of G. morsitans ; and the Director of Mines, 
Magequece, for his kind loan of the Geological Survey Report. Mr. W. M. Longden, 
who spent many months with the Chief Gungunyana, has kindly sent me many 
interesting notes on Manikusa’s and Umzila’s wars. 

Lastly, I have to thank His Excellency the Governor of the Mozambique Com- 
pany’s Territory for his sympathetic interest in the undertaking, and the readiness 
with which he provided for the expenses of the expedition. 


{J.—NATURE OF THE COUNTRY INVESTIGATED. 

The country investigated lies chiefly in the Mossurise but partly in the Moribane 
district, and in the main between the Buzi and Lusitu Rivers. East of the latter 
river's great southward bend it is bounded by the southern termination of the 
splendid Chimanimani massif with peaks of over 8,000 feet, and by that of the 
Mabiti highlands. Its most promiment physical feature is the straight, high, 
narrow Sitatonga Ridge that, pointing to the magnetic north, cuts at right angles 
to the two rivers and effectively shuts off the lowland plains east of it from the 
foot-hill and valley country to its west. The latter is divided further by two rivers, 
the Mtshanedzi and the Puizisi, which flow out from the direction of the political 
border, but turn off, the former to the Buzi, the latter to the Lusitu, shortly before 
reaching the Sitatongas. The valleys are low and broad, as is that flanking the 
Sitatongas on that side, and it was these valleys that were the site of the guard-area 
employed by Umzila for the protection of the enclosed country—a_ successful 
measure against fly which I shall describe below. 

Between the Lusitu and Puizisi, between the latter and the Mtshanedzi, and 
again between the Mtshanedzi and the Buzi, are three hill systems, the first two 
backed by and rising strongly to the highlands of the British frontier (see Map), 
the other consisting of independent second-rate heights rising from broken country 
that falls away gradually to the eastward-flowing Buzi. 

The elevation varies from well over 4,000 feet (1,220 metres) on Mount Umtareni 
and. the Sitatonga crests, and 3,700 at Spungabera, to 2,000 and 1,000 feet and less 
in the lower valleys between the Sitatongas and the British border. East of the 
Sitatongas the general elevation would be nearer 500 to 600 feet. Indirectly at any 
rate, elevation is an important factor here in relation to the permanent distribution 
of tsetse. 

I had no means of taking altitudes, but the following aneroid readings, taken 
by him in 1917, have very kindly been given me by Mr. R. W. Jack. 


Inyamadzi River, on British border av .. 2,200 feet ( 679 metres) 
Lusitu River, near British border by. ~ i 49300), sat eR ipeeer) 
Lusitu, near Haron Junebion,; <4 1,100.55 hin Bee cena) 
Mossurise River on Spungabera-Masanjena Road 950 ng) “adel a) 
Gogoyo, Dysart Concession ce oe L200... a roa) 
Mtobe’ s Kraal Ke Es ys os about LOOT Aes tomate) sa 
Matusi’s Kraal sh As igh pt en eee eee) 
Spungabera .. vel a ie Ata elietay 4h kala 
Jersey homestead .. bs 3 ya Se BOOM STR O MBBS hp) 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 319 


The last figure, though obtained in British territory, gives an idea of the higher 
altitudes in the lower part of Gwenzi’s country, about the Chinyika River south of 
Spungabera. 


II].—WoopbInG OF THE District ANALYSED.* 


A classification of woodland types in relation to tsetse would properly be founded 
on the degree to which and the length of period during which they lose their leaves 
in winter. It so happens that the division between evergreen (or with high, dense, 
heavy undergrowth and lianas) and deciduous coincides here in the main with 
the natural division of the wooding into two principal types. 

These are :—(1) primary forest, pyrophobic or fire-fearmg, dominated either 
by evergreen or deciduous trees ; (2) secondary, pyrophytic or fire-supporting 
forest of various sub-types. The latter replaces the former where that is burnt 
out. Bark adaptations enable it to withstand fires, and it is burnt through annually. 
Where it so thickens up as to reduce the grass growth, and consequently the severity 
of the fires, it tends to be re-invaded by the primary forest. A large proportion 
of the finer rubber forest in the area investigated consisted of secondary forest 
in course of re-invasion, and this harboured the large tsetse (Glossina brevipalpis). 

Primary forest (Pl. ix; Pl. xii, fig. 2) consists of lofty, densely growing trees 
that support many woody lianas and shade lower tiers of evergreen shrubs and 
small trees, with a carpet and fringe that do not readily burn. Primary forest of 
the “rain-forest” type exists in the highlands, mostly in small patches at 
Spungabera and elsewhere, and is characterised by such splendid trees as 
Khaya nyasica (Kast African mahogany or mubaba), Chrysophyllum fulvum (large 
muchanja) and Piptadenia buchanant (umfomoti). The last-named species 
dominates most forest of this type seen in the Lusitu-Sitatonga rubber country. 
It gives it almost the character of “‘ monsoon ”’ forest, for it differs from the others 
in being regularly deciduous ; but the period of leaflessness is not very long, and 
the lianas and high close-packed evergreen shrubs (Pl. xii, fig. 2) continue to 
afford the requisite shade for the forest fly, G. brevipalpis, and to favour its 
activities at all hours of the day. 

There is much striking evidence in favour of the view that primary forest once 
covered this country, and that during very many thousands of years past it has 
gradually been reduced to its present small dimensions through the agency of burning 
and cultivating man. From this and from the analogy of the distribution of our 
other forest animals, quite apart from the results of the movements of game, would 
follow the comforting deduction that primary forests that to-day lack the large 
tsetse possess some advantage, elevational or otherwise, in regard to it, and are 
likely therefore never to harbour it for long. Such forests are those of Spungabera, 
Mount Silinda and the border generally. 

Apparent primary forest here has three sub-types, “‘ mountain,” “ heavy’ and 
“ravine.” The first does not concern us; the second (PI. ix) is characterised by 
the shrubs, Conopharyngia usambarensis and Dracaena fragrans, and the liana, 


‘< 


*These descriptions’of woodland types are necessarily greatly condensed, but their 
recognition will be facilitated by reference to the accompanying illustrations. 


320 Cc. F. M. SWYNNERTON. 


Landolphia swynnertoni; and the third (specially common in the area investigated ) 
is often dominated by the Piptadenra, by the shrubs, Craterispermum laurinum 
and Dracaena gazensis, and the liana, Landolphia kikw (cf. Pl. xii, fig. 2). 
It seems to be the most important in relation to G. brevipalpis, though I have 
also taken this fly in some numbers in the heavier type. 


A river-bank association that is worth mentioning specially owing to its ubiquity 
is that which consists mainly of tall, clean-stemmed trees of mowana (Adina micro- 
cephala) and mubaba (Khaya nyasica) growing on the immediate edge of streams 
and, so long as the moisture supply remains ample, resisting the action of the fires. 
Except where it has been associated with lower woody growth, | have not found 
it to harbour fly. However, it frequently is thus associated. 


Secondary formations—deciduous in varying degrees—present several very 
clearly defined sub-types varying from savannah to close wooding, and each 
somewhat closely confined to a particular geological formation. With ‘‘ wooded 
pasture ” of one or other of these sub-types it covers the bulk of the country. 


> 


The local ‘‘ formations” are as follows :— 

(1). Very highly deciduous types, such as Ptlerocarpus sericeus (mumbhungu), 
Pterocarpus angolensis (bloodwood or mubvangazi)—the earliest shedder and latest 
reclother of all (Pl. x, fig. 1)—Acacia natalitia (ihlofunga), A. caffra (iguwha), 
Bridelia micrantha (mushungunu) and others. Considerable stretches of country 
are covered almost exclusively with bush of these species—open or dense, pure 
or mixed. Fly spreads in during the rains, but I failed completely to find tsetse 
in wooding purely of this type during the leafless period even in such shade as there 
was, and as it is quite unlikely that they can exist there then, it follows that such 
areas are cleared each year for from one or two to many months at a time. The 
country carrying these more sensitive types of wooding is characterised by good 
soil and permanent streams, and much of it by high elevation. It is particularly 
suited to settlement. 


(2). Lowland bush savannah (Pl. x, fig. 2), comprising many tree species, 
but with the mupoza (Combretum near leonense) dominant on the drier parts and 
the musekesa (Bauhinia reticulata) on those that are wetter in the rains. ‘The 
bush is mostly sparse and, the fires being early and the trees and shrubs low, 
the shade is destroyed at an early date in a normal year. Rare scattered fly, 
G. morsitans and pallidipes, was to be found in this type of country up to the time 
of burning, that is in 1918 the end of July, but permanent fly was near by in 
a more suitable type of wooding—Brachystegia, with vleis. I found no trace of 
the existence of permanent breeding centres in this type of country—the basalt. 


(3). Brachystegia wooding, here known as “tondo-bush” or “itondo” and 
evidently identical with what Jack calls “ gusu.” This again contains a variety 
of trees and shrubs, but different species of Brachystegia dominate, such as B. 
globiflora, the mutondo (which gives its native name to the type of wooding), and 
the mutsatsas, B. randii, B. bragaet and B. spicaeformis, the trees of the red spring 
foliage. Uapaca also—U. kirkiana (munjanje) especially in the highlands, 
U. sansibarica (mutongoro) below—is common and in many places forms 
pure wooding. Diplorhynchus mossambicensis (mutowa), with its weeping 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 321 


foliage, creviced stem and branches and furrowed, protectively coloured bark, is 
a particularly useful tree to the tsetse and is abundant in Brachystegia bush at 
low elevations. It is the foreground tree in PI. xvi, fig. 1. 


Tondo-bush is the tsetse bush par excellence. Brachystegia-Uapaca wooding 
in our present connection is divisible into three varieties, two of which carry each 
its own tsetse. It is divisible into (a) high “‘itondo,” characterized by the greater 
size of the trees and generally denser wooding and forming fine “ savannah ”’ forest 
(Pl. xvi, fig. 1); (6) lower, more open “itondo,” still mostly savannah forest, 
but poorer and intermixed with more definite savannah (PI. xv, fig. 1). It is 
(with other conditions correct) the special haunt of G. morsitans. The former, 
finer variety is divisible further into (1) clean itondo, devoid of woody undergrowth 
and of the large fly (Pl. xi, fig. 1), and (2) itondo with woody undergrowth, 
inhabited by G. brevipalpis (Pl. xu, fig. 2). 

This undergrowth, usually patchy, consists for the most part of saplings of the 
trees themselves, often in close-packed clumps (Pl. xu, fig. 2; Pl. xvi, fig, 1), 
and of large scattered shrubs such as the umtunduluku (Ximenia americana) and 
the chigwenderes (Bauhinia galpina and B. petersiana); but in parts of the high 
itondo may be found numerous clumps of “ravine” type primary forest 
undergrowth, extensive or small, of bird and baboon-carried genera that have made 
a start and maintained themselves where the ill-regulated fires have enabled them 
to do so. PI. xiv, fig. 1, gives some idea of their nature. 


Two highly important points about itondo in relation to tsetses are these :— 
(a) The grass-growth in general below high tondo-bush is sparser than in some of our 
other secondary sub-types and does not burn really effectively till late in the season ; 
(6) itondo as a whole, under our local conditions of rainfall, tends to retain its 
leaves through the winter better than any other of our secondary formations. The 
mutsatsas (Brachystegia bragaei, etc.) have often regained their leaves before the 
associated mutondo trees (B. globiflora) have lost theirs, and the Uapacas only 
lose theirs (and that not completely) when nearly all the other species are busy 
regaining their leaves. This tendency to keep in leaf without some s!ight 
interregnum is less marked at high elevations, in particularly cold and dry 
winters, and in the drier parts of the lowland areas; but on the Buzi, and in the 
“ Oblong” (see Map) generally, it was sufficient in 1918 to carry G. brevipalms 
in numbers everywhere through the winter, and it is probable that in most if not 
all years it would do so in most parts of this piece of wooding. 


(4). Dense secondary forest includes especially (a) the very fine type of wooding 
dominated» by musara and musunganyemba (Milletia stuhlmanni and Pteleopsis 
myrtifolia)—definite “‘ monsoon forest’ ; and (6) a more mixed type. In places, 
owing often to the coppicing effect of unthorough native cultivation with the good 
seed-bed offered by old gardens to seeds from the surrounding bush, and to the 
relative exclusion of fire consequent on the absence of grass, the ordinary trees of the 
savannah form more or less dense thickets (Pl. xvii, fig. 1). Eventually trees and 
shrubs that are normally found on the outskirts of primary forest introduce them- 
selves and finally dominate. Such are the munjerenshe (Albizzia fastigiata chirin- 
densis), the large umkadhlo (Rauwwolfia inebrians) and the fine shrubs Vernonia 


322 C. F. M. SWYNNERTON. 


podocoma (mudambasese) and Brachylaena rhodesiana (ipahla). The annually 
burning grasses tend to disappear, the live carpet may become sparse or it may 
become composed in part of the non-burning grasses that occur in the primary 
forest of the same locality. Clumps and stretches of the handsome ribbon-like 
izeya grass (Setaria mauritiana), beloved of bufialos, also of Amomum (good shade 
for brevipalpis) are exceedingly common, and thickets occur of such tall, more or 
less herbaceous plants as Isoglossa mossambicensis and the unpleasantly smelling 
Mellera lobulata. Woody undergrowth is usually more or less scarce excepting 
when the wooding is under invasion by primary elements. 

In the simple coppice stages (Pl. xvu, fig. 1) these woods are highly attractive to 
G. pallidipes ; in the older stages where the overwood is tall and heavy, as on the 
Sitatonga base, and some underwood is present, they shelter G. brevipalpis. In 
the Mtshanezi-Puizisi area especially a large proportion of the groves of this type 
are of relatively low growth and are characterized by the presence of the very 
beautiful climbing fern, Lygodiwm subalatum, which carpets the ground and curtains 
the trunks up to a considerable height, affording low shelter and additional shade 
where the shrub growth is poor. This variety of the type is perhaps more commonly 
frequented by pallidipes than by brevipalpis. “ Dense mixed secondary ”’ occurs 
here chiefly on the dolerite and, like Mulletva bush, tends readily to be replaced 
by primary elements (which are more suitable to brevipalpis) where it is in contact 
with them. Thus Albizzia chirindensis and Piptadenia buchanani (umfomoti) 
are at present sharing the dominance of the Mafusi rubber forest, and the shrub 
growth shaded by them is already largely primary. The musara (Muilletia) is rather 
specially deciduous, and I have found relatively little dry season fly in this variety 
of dense secondary excepting where it was under incipient primary invasion. 

(5). Bauhima and Erythroxylon-Landolphia thickets. Two species of chigwendere 
(Bauhinia galpiom and B. petersiana), large and densely foliaged shrubs that lose 
their butterfly-shaped leaves more or less Jate in the season, occur from the British 
border to the Sitatonga Hills. B. petersvana continues to be present east of the 
hills. They are found in every type of secondary wooding, especially on ant-heaps, 
but in places they form almost pure coppice of their own. When well-grown and 
in leaf, their coppice is more attractive to G. pallidipes. Where, however, the 
Bauhinias occur isolated under higher shade, G. brevipalpis is attracted and rests 
under their coils—for under these conditions they tend to assume the habit of 
lianas. | 

Low dense thickets of Hrythroxylon emarginatum, Pleuroridgea zanguebarica 
and other erect shrubs, with a few rubber vines, have invaded (for the most part) 
Pterocarpus wooding at high elevations on Mount Umtareni. I found no fly there, 
for reasons probably of elevation and also of situation ; for between this and the 
permanent fly-bush (Brachystegia) lower down the slopes lay a belt of highly 
deciduous trees (formation 1, above). Lowland rubber forest (Pl. xi) is not 
essentially different. It also contains much of the Erythroxylon, but the invasion of 
Landolphia and other lianas has progressed further than on Umtareni and here fills 
the thickets. The trees of the highly deciduous and other (Brachystegia) types that 
have been invaded still in many places stand out above the new shrub growth, 
just as on Umtareni. I refer particularly to the Madanda forests, which I did not 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 323 


visit on this trip, but which (from what I saw of them in 1906-7) I should quite 
suspect of being capable of harbouring G. pallidipes and, in many places, 
G. brevipalprs. 


TV.—F actors INFLUENCING LEAF-FALL. 


I have described above the relative deciduousness of our various types of bush. 
We have seen that primary forest (Pl. ix), and bush which it is seriously invading, 
is never leafless enough here to discourage its special fly—brevipalpis; and that 
of the secondary types, Brachystegia wooding (PI. xvi, fig. 1) and particular varieties 
of the “dense secondary’ formation tend to retain in this district sufficient 
leaf to carry fly through the year. 

However, within any particular type of wooding actual leaf-fall is hastened 
locally, and the gaining of the leaf correspondingly deferred, by several factors. It 
is true that some of the differences are apparently individual, though unseen root 
disabilities might account for them. It is at.any rate of frequent occurrence to 
see two trees of the same species and size (e.g. of Pterocarpus) standing together, 
one of them leafless the other not. Ficus subcalcarata (the mutowetowe fig, an 
isolated large specimen of which will sometimes harbour fly) is an extraordinarily 
erratic leaf-shedder. But general rules are apparent also. 

(1). Season affects leaf-fall. After the abnormally heavy rains of 1917-1918 
(2,514-5 mm. at Spungabera) leaf-fall generally took place, I should say, quite 
three weeks later than usual, where frost was absent. A low rainfall (such as that 
of 1913), or early and severe cold, hastens the phenomenon. It must be remembered 
further that rainfall, hence leaf-fall, varies with locality, particularly in relation 
to the mountain ranges. The rainfall appears to be distinctly lower east of the 
Sitatongas than on them or even west of them, and is lower here again than on 
the hills of Spungabera, the “ Jihu”’ (Gwenzi’s) and the British border. 

(2). Proximity of moisture in the soil affects leaf-fall in two opposite ways. 
On damp low-lying ground liable to frost the leaf-fall is hastened, and even where 
there is no actual frost, trees bordering on ill-drained, cold, swampy ground tend 
annually to lose their leaves earlier then those of the same species that stand back 
on well-drained soil. But trees standing on warm, damp soil lose their leaves 
later than the latter. These differences are to be clearly seen in fly-harbouring 
Brachystegia woods, but the particular illustration I shall quote is concerned with 
some scattered wooding of Pterocarpus angolensis (mubvangazi—PI. x, fig. 1) that 
is under my continuous observation. Here not only is there, in May, an increased 
leaflessness as one approaches the cold, damp lower valleys, but of the trees within 
the latter, those with their roots raised on ant-heaps are practically in full leaf 
when those on the general ground-level are completely leafless. 

(3). Differences in the amount of humus present has appeared to account for 
a difference in leaf-fall between woods of the same species—e.g. of Pterocarpus 
sericeus (mumbhungu) on the dolerite—and it is to be noted that ant-heaps exert 
an influence in this direction also. It is probably again mainly a matter of relative 
moisture retention. 7 

(4). Geological formation affects leaf-fall once more on the same lines. I have 
noticed that Pterocarpus sericeus loses its leaves a good deal later on the fertile 


324 C. F. M. SWYNNERTON. 


(and moisture retaining) upland dolerite than on the sandstone-shale formation 
at the same elevation. The Brachysteqia-Burkea groves of the far drier, deeply 
cracking, compact basalt soils of the plains, effectively drained by the low-flowing 
Buzi and highly insolated, were losing their leaves earlier than those of the granite, 
especially on stony outcrops ; while similar wooding on a quartzite strip on the 
Buzi, at the same elevation and distance from the river and only half a mile away, 
was far later than the latter, as was the Brachysteqia wooding (doubtless with better 
rainfall, however) of the sedimentary area of the higher Buzi. 

(5). Altitude, other things being equal, makes a difference. Travelling between 
Maruma and the escarpment at Makwiana’s kraal on 11th August I saw no Albizzia 
(munjerenshe) or other secondary wooding that was not completely leafless. On 
approaching the foot of the descent I began to find some in leaf—and at once found 
fly, both brevipalpis and pallidipes. I have come across many other examples 
both in our individual tree-species and in our woodland types that were thus remi- 
niscent of the instance of the plane tree—simultaneously leafless at Innsbruck, 
yellow-leafed at Garda, dark green at Palermo and in parts of Greece evergreen ; 
and I have little doubt that latitude must also count for much in this connection. 

Altogether it is certain that no unqualified statement disregarding rainfall, soil- 
moisture, frosts, elevation, latitude and exceptional seasons can be made for parti- 
cular types of wooding as regards the period for which they will be without leaves. 
and inhospitable to tsetse. It is, I think, equally certain that an exhaustive study 
of these factors may be of great assistance not merely in the explanation of local 
outbreaks, but in the actual control of the fly. Certainly no investigator can at 
present lay down the law for another area than his own as to the behaviour of 
particular types of wooding and, therefore, for the seasonal behaviour there of the 
fly. 

A point that cannot be too greatly emphasised is that, if we are to fight the 
fly economically as well as effectively, we must base our measures not on the normal 
but on the exceptional season. I do not refer to the season that is so exceptional 
as to occur only once in 50 years—as the great droughts of 1860 (approximately) 
and 1913; nor to cases in which haste is vital ; but, let us say, to the driest or 
coldest season in ten. 

The effect of the factors that I have mentioned above may be seen even when 
leaf-fall has been brought about artificially, as by a fire. 


V.—EFFECT OF THE ANNUAL GRASS-FIRES. 


These very commonly anticipate leaf-fall. Except in the loftier bush, and to 
some extent there, they scorch and wither even the higher leaves and defoliate the 
saplings and shrubs, destroying the shade. The very factors, local and seasonal, 
that accelerate or retard the date of the normal leaf-fall hasten or postpone that 
of the fires also. In 1918—except on the uplands where the frosts were considerable 
and dried the grass—the earlier fires were a month or more later than usual ; in 
1913 they were exceptionally early (after a deficient rainfall) and provided some 
excellent instances of what might be accomplished by severer annual fires towards 
the destruction of tsetse-haunted bush. The Buzi flats with their cracking basalt 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 325 


soil (Pl. x, fig. 2) are stated always to burn earlier than the granite gneiss near 
them (PI. xiv, fig. 2; Pl. xv, fig. 1) with its impervious underlying bed, and the 
sandstone-shale areas of the uplands west of Spungabera definitely do burn 
considerably earlier than the dolerite beside them. The type of bush also influences 
the date of burning. High canopy-forming Brachystegia, such as occurs in 
Mossurise on the upper Buzi, burns effectively much later than the same bush 
where it is lower and more broken, and some types (as high dense secondary, 
including Muilletia) burn, if at all, late yet tamely. 

It is stated that under the Zulus burning was the subject of regulation for hunting 
purposes, a late, thorough burn being aimed at and usually achieved. Under the 
white man everyone burns when he pleases. It thus comes about that the first 
fires take place when the grass is but half ready to burn and there is relatively little 
recent leaf-litter on the ground. They attract but little wind to swirl the flames 
into the tree-tops and the thickets, and they are very small in extent, so that the 
flies driven from them find ample refuge all round. When later fires take place— 
at various dates—the areas burned earlier are already becoming fit for the reception 
of the fly and the latter suffers no inconvenience. A still worse effect of unregulated 
burning, from the point of view of tsetse control, is the encouragement given by 
too early burning to the wooding and particularly to the formation of the thickets 
en which G. brevipalpis rather specially depends (Pl. xvu, fig. 2). Late burning 
on the other hand finds the grass and fallen leaves at their driest and most 
abundant, and is so extensive as to draw in great wind. It achieves much actual 
destruction of young growth, and a temporary destruction of much high shade 
as well, at a hot, dry time that must be relatively critical for the tsetse. 

A failure to burn, again, may be made the means of destroying thicket growth 
if the next year’s burning is late, owing to the additional amount of inflammable 
material present. The fire no longer stops at the outskirts. This result actually 
followed the temporary effort to keep fires from the rubber forests, and I was shown 
where pieces of these forests (in Umtobi’s country—of the type shown in 
Pl. xii, fig. 2) were completely destroyed in consequence. 


VI.—DistrisuTion oF Rocks, WoopLaANpD Types, Foop ANIMALS AND FLy. 


Geological Formations. 


Geologically, the country investigated is divisible into four clear-cut blocks, 
two of them east of the Sitatongas, two west. The two to their east are a granite- 
gneiss and a basalt formation. The two to their west are (a) a sedimentary formation 
of shale, sandstone and quartzite ; (b) dolerite, breaking through and capping the 
sedimentary formation, which in the dolerite area is visible only in places. 

The granite-qneiss country consists of lowland plains, flat to undulating.* It 
is bounded on the west by the Sitatongas, on the north (I believe) by the Mabiti 
hills across the Lusitu and by their north-easterly continuation, on the south by 
a fairly well demarcated line a few miles south of the Umvuazi and apparently 

*My rocks, identified by Mr. Mennell as mica granite, gneissose granite and Muscovite 

anite or gneiss, Show more or Jess foliation, and the area appears to be that, east of the 


Sitatongas, described by Theile and Wilson as metamorphic. The general character- 
istics are those of the granite. 


326 C. F. M. SWYNNERTON. 


following the same general direction as that river. The surface soil is a coarse, 
loose, whitish sand, and numerous vleis, still wet in the dry season, indicate the 
presence below of the usual impermeable layer. 


The basalt plains (PI. x, fig. 2) adjoin the granite-gneiss on its south. They cross 
the Buzi and cover a great extent of country to the south of it. The ground, reddish 
where the rock is exposed, browner or blacker in the interleaved alluvial patches, 
dries rapidly after the rains and shrinks and gapes. In the country actually 
investigated by me it is possible that the Buzi, flowing between fairly high banks, 
determines the height of the water-table. 


The sedimentary area—or rather the area where this stratum is continuously ae 
for it underlies both the basalt and the dolerite—fills the great oblong tract enclosed 
between the Mtshanedzi and Buzi rivers (see Map). A broad strip of the same 
formation runs thence up the Buzi to the British border. It is hilly throughout, 
the soils are mostly compact, and the rock frequently outcrops. Except in the strip 
just referred to, it is not well supplied with permanent streams. 


The sub-ophitic dolerite, as dyke or cap, dominates largely between the Mtshanedzi 
and the Lusitu, and especially at and south of Spungabera in Gwenzi’s country. 
It is accompanied by a red, rather clayey, yet well drained and highly fertile, typical 
“trap” soil, and abounds in excellent permanent streams. 

The Sitatongas themselves are of quartzite, flanked by schists, trap, ete. ; 
apparently (Thiele and Wilson) the schists dominate to the north of them, in the 
area traversed by my native expedition. 


Distribution of Woodland Types. 

The granite-gneiss carries open (mixed) Brachystegia bush usually of a rather 
poor type (Pl. xiv, fig. 2). Its vleis are nearly bare of wooding, except for the 
fine trees on the ant-heaps that stud both vleis and dry land. The finer vleis in 
particular are lined by a short green sedge (Fuirena). The male tsetses are found 
in rather special connection with this. 

The basalt, as might be expected from the above description, tends to carry a very 
poor type of bushy (Pl. x, fig. 2), mixed, but commonly more or less stunted and 
very open woods—the “‘ lowland bush savannah”’ of my classification (p. 320). Thorn 
groves occur in places; elsewhere, on exposed or nearly exposed sedimentary 
rock, Brachystegia patches. The grass, short in some places, is long in others. 
Fires are earlier here than anywhere else. 

The sedimentary rock of the “Oblong” and its northward extension carries 
itondo (Brachystegia-Uapaca forest) of a particularly fine type, with a good deal 
of sapling undergrowth and, in places, patches of invading primary shrub formation 
(v. p. 320 and the figures there referred to). The mutongoro (Uapaca sansibarica) 
often forms close groves beside vleis. The grass growth is seldom dense or very high, 
except in open places (Pl. xiii, fig. 1), but great and annoying burr areas, of four 
semi-herbaceous species of Triumfetta (dzunzu), occur. They dry up and are leafless 
in the winter. 

The dolerite carries (a) wooding of the dense mixed secondary types, largely 
composed of highly deciduous species, and including Bauhinia thickets; (b) open 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 327 


Pterocarpus sericeus (mumbhungu) wooding, highly deciduous. It is also, however, 
the chief seat of the surviving primary forest (Pl. ix; Pl, xin, fig. 2). A high-forest 
area composed of primary forest (largely ravine type), tall, dense, mixed secondary, 
and invasions of the latter by primary forest, covers much of the broad strip of 
country that comprises the Mafusi and Maronga rubber forests, and stretches 
interruptedly from somewhat near the British border at beacons 76-77 eastwards 
to the Musapa, crossing the Lusitu on the way. It is roughly bounded on the 
south by the Puizisi, but sends two broken, narrow extensions southwards along 
‘the eastern and western base respectively of the Sitatongas. Between the Puizisi 
and the Mtshanedzi, and again in the Gwenzi country, lighter dense secondary types 
dominate, copse-like groves (Pl. xvii, fig.1 ), and fern-entwined wooding alternating 
with open, highly deciduous wooding of Pterocarpus sericeus and its Acacia and other 
associates. This more open wooding dominates rather specially at the higher 
elevations, though scattered close Albizzia groves, losing their leaves in the cold 
weather, are also somewhat prominent there. Narrow primary-type thickets 
—fringing forests (Pl. xiv, fig. 1)—often occur along the small streams, and 
harbour fly permanently at the lower elevations. The grass-jungle growth, except 
in the dense wooding, is immense and tangled. The fires are normally more or 
less late and, when postponed further till the grass is dust-dry, they are of 
extraordinary severity. 


Distribution of the Game. 


This again is most simply stated in terms of rock-formation. The lower-lying 
Mtobi-Gogoyo dolerite-dyke country that is bounded by the Mtshanedzi on the 
south, the lower Chikambwe on the west, the Mtobi piece of the Lusitu on the north, 
and the Sitatongas in the east, forms an area that is poorly stocked with game, 
and is surrounded by the heavily stocked areas of the gneiss, basalt, sedimentary 
rock and (latterly) the higher elevation dolerite. Of these well-stocked areas 
the basalt only (Gunye’s, etc.) is somewhat unattractive to the buffalos, though 
they wander into it in small parties in the rains. Their distribution is chain-like, 
the new buffalo colonies of the dolerite connecting occasionally with the herds of 
the sedimentary “ Oblong,” these with the herds of the Umvuazi—Lusitu Brachy- 
stegua-morsitans area (the gneiss), these with further herds in the Moribane district 
(at Muchamba’s, etc.), and these again, it is said, with those of Chermgoma. With 
them are found elands and elephants in numbers, animals that are also rather 
specially attracted by Brachystegia bush, yet have latterly invaded the dolerite 
also. 


Bush-pigs are exceedingly and increasingly abundant on the sedimentary rock and 
the dolerite, far less so on the gneiss and basalt, where they are largely replaced by 
wart-hogs. Herds of sable antelope are distributed through the country generally. 
Lichtenstein hartebeests are present in numbers throughout the two formations 
east of the Sitatongas, also (but much less so) on the sedimentary rock, but not 
at all on the dolerite with its jungly grass-growth. Waterbucks occur in small 
herds on the large rivers. Bushbucks, reedbucks and duikers are present throughout, 
where the conditions are suitable, and little blue duikers (Cephalophus monticola) 
are in some numbers in the dense forests and thickets. 


328 Cc. F. M. SWYNNERTON. 


Baboons are abundant everywhere, especially perhaps in the Sitatongas, in the 
mountains of Spungabera and parts of the British border, and in the dense Lusitu 
wooding. The forest monkey, Cercopithecus albogularis bevrensis, is to be found 
wherever high dense forest is present, and the bush monkey, C. pygerythrus, is 
ubiquitous, in smallish numbers. 


Distribution of other potential Hosts of Tsetses. 


I shall refer sufficiently to man and the smallest mammals below under 
“ Food of the Fly.” But facts to be referred to also, and particularly Lloyd’s 
results from his examinations of the gut-contents of. G. morsitans, show that in 
considering the fly’s dependence on the local game population we cannot leave 
out of account the local bird and reptile population. Guineafowls and francolins, 
the former particularly, both crested and horned (Guttera edouardt and Numida 
mitrata), are everywhere abundant, the crested guineafowl especially frequenting 
dense forest. Of doves, all ground-feeders, two of five common species occur 
in dense forest, three throughout the other woodland types. Each of these two 
main woodland divisions boasts also its own considerable low-searching insectivorous 
bird population and its common touraco. The smaller seed-eaters—waxbills, 
whydahs, weavers, canaries—are a very important item in the bird population 
everywhere except in dense forest, and even there (as I noticed in the Madanda) 
in the open spaces that are covered with grasses that burn. They are nearly 
always found on or near the ground, and commonly in great flocks. They are 
most abundant in the non-forested, heavily seeding dolerite areas—with less 
permanent fly. Quails were seen everywhere in grass-country and much on the 
borders of the morsitans vleis. 

Another bird that haunts every type of woodland and has been recorded as being 
attacked by tsetses (Lloyd) is the ground hornbill (Bucoraz cafer), purely a ground- 
searcher. It walks in small spread-out parties and (in common with the insect- 
tivorous birds) makes up somewhat for any lack of numbers by adopting a definite 
beat, most parts of which it revisits at intervals of a few days. In common with 
them also it probably does the fly more harm than good, for it destroys many snakes 
and the young of ground-nesting birds. Secretary birds and the various storks may 
be classed with it in this respect and are found in rather special association with the 
two formations about to be mentioned. | 

The water-haunting birds are found mainly on the basalt (on the Buzi) and 
granite-gneiss. Jacanas, egrets, bitterns, hammerheads, moorhens, rails, grebes, and 
occasional herons are found at the larger pools of the vleis of the latter formation, 
constantly but in small numbers and in contact with some of the dry season centres 
of G. morsitans. The two largest rivers and such fine series of pools as those east of 
Chibabava boast a far larger water and waterside population, but, so far as I have 
seen, it does not come into really intimate and essential contact anywhere with the 
main tsetse populations. The ground-running birds—the various bustards and 
coursers, dikkops, sandpipers, plovers, etc.—are particularly addicted for many 
spring and summer months to the shorter-grassed and sparingly wooded basalt, and 
certainly also to the granite-gneiss, though I have not seen them there when the grass 
has been recently burnt. 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 329 


The main reptile-population follows closely the two biggest rivers and is not 
sufficiently intimately connected with the big permanent fly-belts to be worth 
discussing. In the fly areas I noticed more snakes, including several puff-adders, 
on the granite-gneiss than elsewhere. Other reptiles were a large arboreal Varanus, 
which occurs in various types of secondary wooding, including Brachystegia, but of 
which I have seen few individuals in all, and some not very abundant Agamid and 
smaller lizards in the Brachystegia areas. 


Distribution of the Fly in relation to the Geological Formation. 


The granite-gneiss, with its open-stemmed Brachystegia covering and frequent 
vleis, is the special home of G. morsitans, which in the dry season is still to be found 
in numbers and breeding actively beside particular vleis. G. pallidipes is also 
plentiful there, and G. brewpalpis is found in small numbers where the shade is 
particularly heavy. 


The basalt, with its sparse stunted bush, was still carrying an extremely thin 
sprinkling of G. morsitans and pallidipes in late July, when burning was barely 
beginning. I am inclined to regard it as, in the main, an overflow area from the 
gneiss, and it is quite likely that any measures that banished fly from the latter 
formation would incidentally banish it from the adjoining basalt. 


The sedimentary rock area, with its fine Brachystegia forest and considerable if 
patchy undergrowth, is apparently more or less heavily infested almost throughout 
with G. brevipalpis. G. pallidipes also occurs, but in small numbers, said to be 
greater in the rains. 


Of the two woodland sections of the dolerite area the Mafusi-Musapa high-forest 
section, with its extensions, appears to be more or less heavily infested throughout, 
at any rate in the dry season, with G. brevipalpis, G. pallidipes also occurring. 
Pupae of G. austent were taken in numbers near the Buzi-Mtshanedzi confluence. 


In the Puizisi-Mtshanedzi area on the other hand G. pallidipes, though in small 
numbers seems rather specially in prominence, perhaps in part owing to the relative 
absence of the bigger fly. The latter does occur here and there in small numbers 
wherever the shade is suitable. Behind the Makwiana escarpment the country 
appeared to be clear of the fly by early August and had probably been so for some 
weeks, though it would still doubtless be possible occasionally to encounter there 
an odd straggler carried from below, particularly on the Chikambwe’s tributaries, 
where leaf was present. At the lower elevations, where also the bush of the dolerite 
retained more leaf, fly was still met with ; but judging from the observations that 
Dr. Lawrence has most kindly carried out for me since my departure, it would seem 
that it must become specially scarce there also towards the close of the dry season, 
though not extinct. It is in part a matter of leaf-fall, for at his station (Gogoyo) 
pallidipes showed itself once more in some numbers with other biting flies when the 
leaves were well in evidence again in the hot weather shortly preceding the rains. 
Glossina brevipalpis was taken in small numbers here and was seen by me also on the 
Buzi below Spungabera. 


An expedition sent by me to the Chinyika in late September failed to take any 
tsetses, though it brought back numbers of other biting flies. 


330 C. F. M. SWYNNERTON. 


There are three great permanent fly-areas in the country investigated :—(1) The 
morsitans-pallidipes area of the granite-gneiss ; (2) the brevipalpis area of the 
Lusitu rubber forests ; (3) the brevipalpis area of the “‘ Oblong.” 

Between the last two is sandwiched an area of more deciduous foliage. Out of 
the higher parts of this such fly as has got there is ousted in the cold weather by the 
leaf-fall, and in its lower parts the fly becomes scarcer and more localised before the 
rains, reappearing with the leaf in October. These last remarks refer to G. brevipal- 
prs and pallidipes, the latter fly occurring throughout the area. 

Finally, there is a narrow strip along some of the frontier, widest at Spungabera 
and on the hill-mass enclosed by the Puizisi-Muchamba confluence, that may be 
regarded as quite fly-free, excepting in so far as travelling parties of male flies may 
on occasions be carried into it during the rains and early dry season by buftalos, ete. 
Nearly all the country on the British side of the frontier may be regarded as falling 
within this category. 

G. austent was taken only near the junction of the Buzi and Mtshanedzi. The 
type of wooding in which it occurred is not uncommon on lowland streams, and 
there can be little doubt that further investigation will show it to have a more 
extended local range. 

More generally it may be concluded that each fly is confined closely to particular 
woodland types, which, in turn, are confined to particular geological formations, 
the conditions that suit pallidipes being the most widely distributed. But the 
distribution of the permanent fly is by no means thus conterminous with that of 
either the buffalos, the big game generally, or any particular species of mammal, 
though abundant mammalian life of some kind is present in each of the areas, fly- 
carrying or otherwise. Man is relatively scarce (except on the Lusitu), and very 
small mammals are abundant everywhere in the fly area. The dominant bird 
elements of the brevipalpis areas are insectivorous and gallinaceous. The morsitans 
area has in addition a stronger bustard-plover element and, at some of the 
permanent fly centres, a very limited water-bird element, as well (apparently) as a 
somewhat larger reptilian population than the other areas. I may add that 
various moth larvae, numerous, gregarious and often immense, are much more 
in evidence in Brachysteqia areas than elsewhere ; but, for reasons to be stated, 
it is doubtful in spite of Roubaud’s results whether they are of great use to 
the fly. The fact that brevipalprs does not come readily to man makes its 
presence difficult to detect unless one is lucky enough to come across a male queue 
some evening. However, my conclusion that the “ Oblong ”’ was well infested was 
come to after I had seen only two flies. Their hum in flying struck me as being 
identical with a sound I had been hearing continually since I came there. I then 
made a point of disturbing all large flies of any species I saw and made my natives 
bring me for a day or two all they could catch for release close to me, but I failed 
completely to match the sound, which I therefore concluded to be that of G. brew- 
palpis. I continued to hear it, often on the part of large flies which I could not then 
identify visually that came out to us and turned back, evidently rejecting us, and of 
flies that flew away from the shady sides of tree-trunks when these were beaten. 
An observation on pigs of a day or two later and subsequent results from cattle 
abundantly justified the conclusion. 


THE TSETSE PROBLEM IN NORTH MOSSURISE. aol 


Vil.—Umnzita’s EXPERIMENTS IN CLOSE SETTLEMENT. 


Earlier History of the District. 


About ninety years ago, following immediately on the invasion of Bulindhlela or 
lower Gazaland (the Delagoa-Inyambane country) by the Zulu chief Manikusa, what 
are now the Mossurise and South Melsetter districts became the scene of three 
invasions. First came that of “the Bulindhlela chiefs, who, though defeated by 
Manikusa, refused to submit to him. . . . They retired in a north-westerly direction, 
over-ran this country, seized all cattle (then plentiful) and later proceeded north to 
Matoko’s.” Secondly, these fugitives were “ followed by Unéaba (alias Mosani) as 
far as Chiamiti—now the farm Schaapplaats in South Melsetter. Here Unéaba and 
his people settled.”” Unéaba was a brother of Manikusa’s father, Isigode, and he 
and his people were part of the Zulu invading force under Manikusa. 

““Unéaba became unpopular here, one of the principal grievances being that he 
would not allow members of his crack regiment, the Amakanda, to marry until their 
heads were grey. Representations were made to Manikusa and he was induced 
to come up here. Fighting occurred and Uncaba, forsaken by his best fighting 
regiment (the Amakanda) retired with a following up the Sabi valley, and 
subsequently in a north-westerly direction” (letter from Mr. W. M. Longden). 
I do not know whether the view is correct that this “ impi” founded the 
“ Angoni” tribe near Lake Nyasa. 

Manikusa (who returned to lower Gazaland) died in 1861. His second son, 
Umzila, disputed the claim of his elder brother, Mawewe, to the throne and, at first 
defeated by him, went north and settled in succession at Umpombo’s on the Buzi, 
and Dongonda north of the Lusitu (see Map). Mawewe eventually followed him 
thither, but was defeated with the aid of the Portuguese, and Umzila took possession 
of lower Gazaland. He himself remained at his northern capital, which he shifted 
more than once. From Dongonda he moved to Gandwa, on what is now the writer’s 
farm “ Gungunyana ”’ in South Melsetter, and thence, later, to the neighbourhood 
of the Chinyika River in Gwenzi’s country, south of Spungabera. Here he made 
his longest stay, eventually moving to Mwayamuhle’s, just south of the Mossurise 
in Garawha’s country ; and here he died. 

His son Gungunyana shifted the headquarters across the Mossurise to the foot 
of Mount Singuno, but later, in about 1889, he returned to the old southern capital 
Bileni, near Lourengo Marques. The population of Mossurise and South Melsetter 
was forcibly collected and taken thither. Soon after—in 1895—the Portuguese 
Government had to bring Gungunyana to account, and with his capture in the 
following January (Theale), the Zulu native domination ended. Since then the 
Mossurise natives have gradually drifted back from Bileni to their old homes, but 
it is stated on all hands that only a small proportion actually has come back, and 
that the population of the district does not at all approach its former size. 

In 1893, the land on the British side of the border was occupied and settled by 
Europeans, that on the Portuguese side, in part depopulated of natives as I have 
stated, remaining wild to this day. In 1896-7 the rinderpest swept through the 
country and, missing small areas here and there, decimated not merely cattle, 
but in particular buffalos, kudus and elands. 

(737) B 


oon C. F. M. SWYNNERTON. 


Fly and Cattle before Umzila’s Time (say 1820-1861). 

There can be little doubt that ever since the arrival of the Bantus particular 
pieces of country have been subject to local concentrations of population, and that 
where other circumstances have been sufficiently favourable, these concentrations 
have resulted, while they lasted, in the opening-up to cattle-farming of the piece 
of country involved. 

The “ old people ” had told my various native informants that previous to Unéaba’s 
massacres of about 1830, farming went on successfully even close to the Sitatongas, 
where they had shown them the old cattle pits. The country was very densely 
settled. The chiefs Mafusi and Mtobe have both told me that at that time cattle 
were kept in those portions of their countries in which all subsequent attempts 
to keep them have failed, and Gunye has stated that it had been handed down that 
in the days before Uncaba cattle-farming, accompanying a concentrated population, 
took place even in his country, but that many used to die. But when Umzila 
came here (in 1861) the whole country had reverted to wooding and game as the 
result of the depopulation arising from Unéaba’s massacres. It was much as it is 
now ; fly had become plentiful, and the mountains of the present political border 
were the boundary, then as now, between the fly and such cattle as existed. 


Umzila’s Experiments (1861-1889). | 

Both when at Umpombo’s, and later at Dongonda, Umzila made a determined 
attempt to keep cattle, but in each case he failed. The concentration of population 
was not so great as in his later attempts ; it was short-lived, and close by was a great 
belt of G. morsitans—a fly which follows man. 

The Zulus, it may be mentioned here, knew the fly well, and the disease caused 
by it, and they regarded the proximity of game as dangerous for cattle. But why 
did they place their cattle amongst the fly? They said, “this whole country is 
full of it—where shall we put our cattle ?” However, cattle had always thriven 
in the open hilly country which is now South Melsetter; they were being captured 
at this time (after Umzila’s defeat of his brother) in great numbers in raids on the 
Mashonas, and Umzila at last sent them all to Gandwa. Later he said ‘“‘ I cannot 
live away from my cattle,” and it was thus he left Dongonda. 

From Gandwa, Umzila sent an order to sondela enkosini (draw near to the King). 
Thereupon an immense compulsory movement of the population took place. The 
country to the east of the Sitatonga Hills, particularly in and south of Gunye’s, 
was at that time more fully populated than that to their west, but almost the whole 
of this population was deported, territorial chieftains and all, to the lower parts 
of the tract between the Sitatongas and the present British border, to Spungabera 
and Gwenzi’s country, and (with Umzila’s further movements) to the Umswirizwi 
(Mossurise) valley, to Zinyumbo’s Hills and Chimbiya’s, and even eventually to 
the Sabi east of this area. The chief Gunye told me that he himself was compelled 
to settle at Chimbiya’s. 


The Effect on the Wooding. 
Every one of my informants has described most graphically the result of this 
concentration. The bush simply disappeared and the country became bare, except 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 333 
for the numberless native villages and a continuity of native gardens. This complete 
clearing was confined to the lower-lying areas. The Puizisi-Mtshanedzi hill-mass, 
as Makwiana told me himself, was relatively little affected by the new settlement, 
and the bush was never at all completely cleared. It was, however, surrounded 
on three sides by a broad cleared cordon, and on the other, backed by the highlands. 


In Mtobe’s and Mafusi’s the bush was never fully cleared ; the rubber forests 
already existed and they continued to do so, although, according to Mafusi and. 
others, they then consisted merely of umfomoti wooding (Piptadenia) that was 
chiefly confined to the ravines. In Gungunyana’s time they had already spread 
greatly (under encouragement), and many Landolphia vines existed throughout 
the new areas not yet wrist-thick. They were a valuable asset to the Zulus, who 
traded the rubber for cloth on the coast, and used the latter in turn in the barter 
of cattle. Again, away from the neighbourhood of the Mtshanedzi and the Buzi 
the ‘“‘ Oblong ”’ remained completely uninhabited and uncleared. 


Zinyumbo’s area, like Gogoyo’s, was very completely cleared—“right to the 
Mwangezi it was gardens only,” as was Gwenzi’s country, the Mossurise valley 
and portions of the Sabi. 


The Effect on the Game. 


Large mammals became very scarce—not merely big game, but pigs and baboons. 
“The Mangoni (Zulus) are killers of everything, men and animals.” Drives with 
nets were organized across the entire country, and game, pigs and baboons were 
thus killed wholesale. If a herd of buffalos was reported subsequently anywhere 
west of the Sitatongas, it was at once hunted ; if pigs appeared in a garden, they 
were at once tracked down to their retreat, and, the people round having been 
called out, were surrounded and killed. Except on its fringes the “ Oblong,” 
then as now, was a great uninhabited game reserve, The game in it was thinned, 
it is true, and was kept well driven within its borders, but there still remained enough 
to attract the Mangoni hunting parties. In the heavily settled areas a few bush- 
bucks, duikers and pigs were still to be found throughout the period. 


The Effect on the Tsetses. 


There were still plenty of int-hesi (tsetse) in the tondo-bush on the granite-gneiss, 
and fly never disappeared in the “ Oblong.”’ In neither place could cattle ever be 
kept. In Mtebe’s country too and the eastern part of Mafusi’s—that is, within 
a short distance of rubber forest—they could never keep cattle. Mafusi told me that 
he remembered the fly there, with the same distribution as now, from the time he 
was a child, except that it has spread with the spread of the forest. At Maronga’s 
and beyond—that is, in the continuation eastward across the Lusitu of the rubber 
forests—cattle could never be kept, though Usele and other Zulus settled there 
and made the attempt. In such places the cattle required for ceremonial purposes 
used to be brought, as needed, from the safe areas. Cattle also continued to fail 
in Gunye’s country and south of the Buzi from the Mwangezi eastwards, though 
persistent and prolonged attempts were made to keep them in this area of sparse 
fly, the losses in some cases being made up time after time from outside. 

(737) B2 


354 C. F. M. SWYNNERTON. 


In Zinyumbo’s hills and even just on the Mwangezi cattle succeeded, as they did 
thence westward through the Mossurise valley and northward to Spungabera. 
This tract was full of cattle. But few deaths took place, and these always along 
its extreme eastern margin, on the borders of the “ Oblong.” 

In the Gogoyo-Makwiana tract cattle were kept right under the Sitatongas both 
at and opposite the Rupisi and from the great bend of the Mtshanedzi to its source, 
also in the hills behind the cleared guard-area between the Mtshanedzi and Puizisi 
and up to and beyond the present British border. Many of the herds were large. 

On the Sabi (present British territory) the results were particularly interesting, 
for this had previously constituted a separate fly-belt, which was eventually almost 
completely wiped out by native cultivation. The rinderpest may have given the 
coup de grace to the surviving remnant or two (recent events render this doubtful), 
but at any rate cattle were already being placed and kept successfully all over the 
old fly area in the seven years between Gungunyana’s departure (with all the cattle 
he could take) and the advent of the rinderpest (W. M. Longden and others). 

When the country was closely settled, cattle were kept successfully in places 
where they had always died before ; and when the settlement was well established, 
they succeeded where in its earlier days they failed, though fluctuations still took 
place with successive shifting of the population. It is true that herds actually 
abutting on the fly still suffered small and occasional losses, as they. are doing to-day 
to a greater extent on the present fly boundary. 


What drove the Fly out ? 


From the failure throughout the 25 years or so of this experiment to keep cattle 
near the Mafusi rubber forests, well settled and well cleared of large mammals 
though they were, it is likely that it was not the mere destruction of its food supply 
that cleared the Zinyumbo, Gwenzi and Puizisi-Mtshanedzi areas of fly. We may 
compare also Mr. Pollard’s statement with regard to the Munshi division of Northern 
Nigeria (Bull. Ent. Res. 111, 1912, p. 221). “It is interesting,” he writes, ‘“‘to note 
that the Munshis are great hunters and that they have practically destroyed all the 
wild game in their district, and yet, in spite of this, the trypanosomiasis of cattle 
and horses is rampant.” 

But the game does not merely feed the fly ; it also carries it, each year, from the 
permanent fly-areas into the more deciduous areas adjoining them as soon as these 
have sufficiently regained their leaves. The heavy settlement of the broad river 
valleys and the base of the Sitatongas, by clearmg the bush, not only rendered 
those areas themselves inhospitable to the fly, but also opposed a barrier to the 
passage of game, and so protected from the fly the large enclosed piece of chiefly 
deciduous bush behind the guard-area. Previous to the establishment of the guard- 
area it had been impossible to keep cattle in the area behind it, as it is now again 
impossible, and the broad guard-area itself, to judge from the descriptions of 
the bush cleared, had probably harboured even more permanent fly then than it 
does to-day. That the fly-population itself was diminished by the Zulu system of 
late burning is also by no means unlikely. 

The case is most instructive and points the way for future measures. It must 
be remembered that the flies concerned—brevipalyis and pallidipes, especially the 


THE TSETSE PROBLEM IN NORTH MOSSURISE. BaD 


former, which is more abundant—are not such great followers of man as is morsilans, 
and that the chance of success in such a case, without wholesale clearing of the 
bush, is greater than where morsitans is concerned. On the Sabi it was different. 
Here the permanent fly-bush itself was destroyed to a far greater extent and the 
fly in most parts exterminated. 


Was the Experiment intentional ? 


These concentrations of population round the King were not primarily fly- 
measures ; they took place whether the country was fly-infested or not. But their 
effect in fly-country was well known to the natives, and I was informed that Umzila 
entertained considerable hopes in relation to the keeping of cattle from his successive 
concentrations at Umpombo’s, at Dongonda, and west of the Sitatongas. Success 
was attributed to the effect on the game rather than to that on the shade, and 
Umazila’s measures against the game were stringent. His intention of colonising 
the “ Oblong” was frustrated by the depredations of lions on the settlements 
placed there (e.g. in the Budu country), and there were difficulties connected with 
water and an immemorial and bloody feud with regard to the Dengaza tract between 
the chieftains Makwiana and Gogoyo;* but he settled its borders very heavily 
and kept the game well driven within them by means of frequent hunts. 


The Effect of the subsequent Depopulation. 

Before Gungunyana carried off the population to Bileni (near Lourengo Marques) 
in about 1889, he had already commenced to protect the game. He had decreed 
az-zale (let them multiply), and game had become more abundant both outside and 
inside the cattle-keeping areas. The guard-areas still opposed its passage into 
and out of these areas and no harm resulted to the cattle. When the population 
left, the game (in the words of my native informants) just “ burst forth” (za- 
dabuka). At the same time the wooding was let loose and soon re-established 
itself throughout the previously settled country. 

In a very few years (by 1896 according to native information) the fly had more 
or less regained its old wet season limit—not many miles east of the present political 
boundary—though the bush capable of supporting it was not then so ubiquitous 
as it is now. I myself lost a beast from fly on “Scott's Hope” (a Portuguese 
farm on the British border) in 1900, and a tsetse was taken south of Spungabera, 
on the tondo-covered hills west of the Buzi (spot shown on map), in the same year 
by Drs. Wilder and Lawrence. 

The fly caused little trouble then, but two things have happened since—a pro- 
gressive invasion of the upland dolerite (a) by the deciduous wooding, and (b)—more 
important still—by game of wandering species. Buffalos and elands especially, 
decimated by the rinderpest, are now extremely abundant, and having spread from 
the sedimentary area into the hills, wander much more frequently than they used 
to into the highlands. It is only in the last three years that bufialos have reached 
the Puizisi country in any numbers. They have there tapped another brevipalpis 
area, and the consequence has already been visible in the destruction of the cattle 
of the Muchamba-Puizisi valleys. 


* Not that this was allowed to continue actively under Umzila. He decreed * There 
must be no more war. We are the only King (Si’nkosi sodwa) ! ”’ 


336 C. F. M. SWYNNERTON. 


VIil.—Tue Foon or tae Fiy. 

Observations by numerous workers elsewhere have shown that tsetses living 
in contact with large reptilian and avian populations very freely utilise the former, 
and perhaps also the latter as food. Gut examinations even of G. morsitans have 
shown in some cases (Lloyd) a remarkable proportion of non-mammalian contents ; 
the same fly has been seen to feed on the bare neck-wattles of an unconfined ground 
hornbill (also Lloyd) ; a case is on record (Ensor) in which a fly, “ almost certainly 
morsitans,’ was observed through glasses trying to get under a perched hawk’s 
wing ; and in the Entebbe laboratory Glossina palpalis “fed readily on captive 
fowls, creeping under the wings to reach the poorly protected parts”’ (Bagshawe). 
It follows from this last and extraordinarily important observation (which 
does not surprise me when I remember that morsitans rather reminded me of Olfersia 
‘in its low poise and clever sidelong movements in response to attempts to catch it) 
that birds are not necessarily protected against tsetses by their armature of feathers, 
though many of them certainly are by their agility and insect-eating habits. Yet 
young birds, whether nestlings or runners both in the nest and for a time after 
leaving it, are defenceless—and abundant just when the fly is abundant. Also 
the fly (as met with on this trip) retires to rest later than do most diurnal birds, 
the latter being thus relatively unprotected from it during a very brief portion 
of each day when the fiy is particularly active—but the bird (it must be admitted) 
less conspicuous. 

Such facts, with the way in which birds, including ground hornbills, keep to 
definite beats, the whole of which they work every few days, and the fact that nearly 
the whole bird-population searches for its food and breeds either usually or very 
commonly within a few feet of the ground, the search being conducted with much 
bustle and conspicuousness on the part of the large combined parties of small birds, 
must be taken into account when we attempt to explain such cases as Umzila’s 
failure to keep cattle near the then much smaller rubber forests (in spite of a great 
hunting population and a strenuous eflort to exterminate game) and when we 
consider the question of destroying game to starve the fly. At the same time, 
so far as small birds are concerned, it seems to me that the fiy’s habit of choosing 
harmoniously coloured surfaces to rest on suggests that its effort will be to evade 
rather than attack them during their active hours and stages of existence. I feel 
aiso that the very striking connection between the fly’s breeding habits and game 
raust be given its full weight. 

My own observations dealt only with mammals. 


Mammals from the smaller Unguiates downwards. 

The natives of the morsitans area were unanimous in their statement that tsetses 
feed freely on baboons, and that “ wherever you find baboons you will also find fly.” 
They gave me many instances in which baboons driven from their gardens had 
left numerous replete flies behind and others in which flies were attracted in numbers 
to baboons that were killed. Simpson’s similar personal observations and native 
statements on the Gambia and Gold Coast will be recalled, as will Lamborn’s 
indirect evidence. Monkeys (Cercopithecus pygerythrus, which lives in lower bush 
than C. albogularis bewensis and raids natives’ gardens) were stated more rarely 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 387 


to have done the same, and—an important statement if true—it was stated that 
replete tsetses were occasionally found where cane-rats (abundant in much of this 
country and very largely diurnal) had been many together. 

Pigs were as universally incriminated as baboons, and here I obtained a quite 
excellent instance myself. In the “Oblong” (east end) in cloudy weather we 
walked right on to four bush-pigs sleeping. In their hasty rush they left the flies 
behind, and these streamed after them in great numbers and with quite a hum. 
We captured nearly twenty that through repletion could scarcely fly ; all but one 
were brevipalpis, the exception being a pallidipes, and more than half were females. 
Austen records both morsitans and brevipalpis as feeding freely on wild pigs and 
quotes Dr. Hearsey’s statement that G. morsitans was seen to settle literally in 
hundreds on the carcase of a wart-hog, behind which animal | also took these flies. 

My indirect evidence of the value of such animals as pigs to the fly was also 
interesting. In a mile-wide patch of primary forest east of the Sitatongas, in 
which I saw much brevipalpis, both native information and a careful search for 
spoor showed that it could have been feeding on nothing but pigs, baboons and 
smaller fry. The same applies to a piece of high, dense, secondary forest on the 
western foot of the hills, into which according to the owner of a kraal on the spot 
(confirmed by the usual search for spoor) no big game had entered for some months. 
It also applied, I am certain, to much of the rubber forest area in which pigs and 
little blue duikers—and these only—are abundant. In one place a length of about 
300 yards for a very great width was continuously turned up by the pigs and looked 
like a hoed plantation. It made me wonder whether one of the forest hogs 
(Hylochaerus) may not occur in these forests. 

Finally, working for over a fortnight round my camp on the Buzi east of Spunga- 
bera, in an area in which game is relatively abundant, I made a special point of 
studying the daily spoor in relation to the distribution of G. brevipalpis. To sum 
up the result, there was a considerable area that I am certain was not entered by 
big game during my stay, or for some days before it. There was a smaller, inner | 
area, immediately round certain kraals, that had probably not been visited by such 
animals for many weeks or months. In genera] the spoor showed much less move- 
ment of the big game now than in the wet season, yet the fly was equally present 
throughout, lurking in all the thickets to attack passing animals. 

The only “ passing animal” that showed a similar ubiquity was the bush-pig, 
and I was convinced from the evidence that the fly was living practically entirely 
on bush-pigs at the time of my visit. Man was not being attacked. My friend 
Mr. G. D. Otterson spent a few days with me here, and declared on leaving that 
the evidence of the fly’s independence of the bigger game in the matter of food 
was a ‘‘complete eye-opener ”’ to him. 

I conclude from these observations that any attempt in north-west Mossurise 
to destroy the fly by starving it in its permanent haunts is doomed to failure if the 
bush-pigs, and perhaps the baboons also, are not destroyed ; and the destruction 
of the pigs in this type of country is not easy. 

Two species of lemur occur in the fly areas. The larger of these, Galago crassi- 
caudatus, is abundant in most of the brevipalpis country and sleeps exposed (but 
perhaps protected by its fur) in the daytime. Bats, including the largish fruit-bat, 


338 C. F. M. SWYNNERTON. 


Epomophorus crypturus, are abundant, and insectivorous species were found sleeping 
in great numbers in hollow trees of Sterculia trvphaca on the ant-heaps of vleis 
frequented by G. morsitans and G. pallidvpes. Bushbucks and duikers are more or 
less common—and hard to exterminate. The little blue duiker (Cephalophus 
monticola), more or less abundant in thickets and dense forest, is less hard to reduce 
by fence-trapping. It has a habit of resting under logs and between tree-buttresses 
that might bring it into additional contact with the fly. It was one of the animals 
least liked by my carnivorous mammals. Of yet smaller mammals two species of 
Tatera came in great numbers to my traps at a spot where male clusters of morsitans 
were present all the time on the grass and females in the wooding (PI. xiv, fig. 2). 
Lloyd has suggested that nocturnal mammals, including Murip4g, are perhaps 
liable to the attack of tsetses owing to the latter’s habit of resting in similar holes. 
T have myself taken morsitans puparia from holes made by animals in a bank, and 
noted that brevipalpis and allidipes rest in holes in trees. Nevertheless, an 
argument exists against this view in the fact that some of the bats at any rate are 
very easily disturbed, and the part played by the small nocturnal mammals, including 
bats, needs far fuller testing than it has received. Species of Arvicanthis, Otomys, 
Myosorez and Petrodromus are largely diurnal, but their small relative size 
suggests both their ready perception and intolerance of the fly and their less ready 
detection by it. ; 

Dassies or rock-rabbits (Procavia) may be of much use to the fly, but very locally 
and only in rocky places. I found them in the coarse quartzite rocks that form 
the crest of the Sitatongas and support Brachystegia; and I spent many hours 
with several workers trying to find wet season tsetse puparia in the humus under 
raised and overhanging rocks, but without success. I searched similarly in the 
Maruma forest, frequented by dassies, but Maruma, I judge, in spite of the outbreak 
of trypanosomiasis there, is seldom if ever reached by the fly. 


The larger Ungulates. 

I have thought it well to give prominence to the evidence favourmg the view 
that the pigs, aided doubtless by the smaller buck and perhaps yet more by the 
baboons, will continue to support a fly population in parts of north Mossurise even 
if the larger game is banished. At the same time it goes without saying that the 
larger the permanent game population the greater will be that of the fly; and I 
obtained abundant indirect evidence that, as things stand now, the bigger ungulates 
present in the fly-areas are taking an important part in the feeding of the tsetse. 
As I shall describe elsewhere, I found numerous batches of puparia—mostly of 
morsitans, a few of brevipalpis—associated with spots where larger mammals had 
been lymg down. Buffalos were incriminated the most frequently, but Lichtenstein 
hartebeests, wart-hogs and other species were concerned. 

My finds of pupae in the morsitans area seemed rather to suggest a preference for 
the buffalo on the part of the fly. I found this definitely asserted by some of the 
natives and denied by others. My guide in the morsitans area, a very observant 
native, was particularly convinced of it. Questioned as to a connection between 
the two animals, he replied that one may find fly where there is no buffalo, but that 
where there is a choice it follows the buffalo. ‘“‘ The buck,” he said, ‘‘are much 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 339 


more restless under its attentions than the buffalo, the hartebeests especially keeping 
up their dance when tsetse are about them; so that the fly can feed more easily 
on the buffalo.”” Neave has already made a suggestion to this effect. 

I shall discuss this subject further under the next heading. Elephants were 
stated by the natives to be attended by tsetses when the latter are numerous, and I 
took a male brevipalpis waiting on a much used elephant path. Roubaud and 
Bouet (referring to G. longipalpis) are both quoted as speaking of a special association 
between tsetses and the elephant and hippopotamus. 


Man—and the Fly’s Preferences. 

Using cattle as I did, I obtained abundant and excellent evidence that G. morsitans 
and G. pallidipes attack these animals more readily than they attack man, that 
G. morsitans attacks man more readily than does G. pallidipes, and that the latter, 
in turn, attacks man far more readily than-does G. brevipalpis. It constantly and 
everywhere happened that the carriers passed through a place without drawing 
fly and that the cattle coming just afterwards attracted many. A dozen to twenty 
carriers are more conspicuous than two or three head of cattle and their scent is 
overpoweringly stronger. Similarly I have seen a fly (pallidipes) on a leaf beside 
the path allow carriers to pass it unmolested and then at once fly out to my donkey. 
Another piece of evidence was afiorded by a fly (brevipalpis) that 1 found perched 
on a projecting slab of shale in the path, facing away from me. I stopped short 
(about two yards away) and had the cattle (immediately behind me) turned into 
the grass. After about two minutes the fly flew back to me and, without alighting, 
returned to its perch. I then had the two cattle immediately brought up, the 
leading ox standing nearly level with me in the path. The fly shortly flew back 
again, first to myself, then, swerving off, to the ox, under which it at once settled 
and began to feed. 

A point that struck me in the morsitans area was that whereas when we were 
digging for puparia there was often delay before a fly attacked us (one fly was actually 
seen watching on a tree-trunk for a time before it came), the cattle were attacked 
quickly. 

Man is attacked by all these flies much more frequently in the rainy season, but 
the increased attacks are probably not more than proportionate to the great increase 
in the number of the flies at that time of year. Dr. Lawrence has sent me records 
of occasional attacks on natives at his homestead at Gogoyo’s even by brevipalpis, 
and as for morsitans (no doubt in company with pallidipes), I was informed by the 
natives across the Sitatongas that they become at times unbearable, forcing every 
man to carry a leafy twig with which he continuously switches his back and shoulders. 
G. pallidipes (it is stated) also becomes a nuisance in places west of these hills, 

Concerning the practical question, “if other sources of food were eliminated, 
could the tsetse still keep going with the aid of man?” it is certain from the 
observations of Lloyd, Maugham, Stephenson-Hamilton, myself and others that 
the tsetses would then attack man much more. Tsetses (as I have seen) constantly 
obtain full feeds from man and escape unscathed and, where the bush comes up 
to a village, so far from avoiding it, morsitans and pallidipes—but not brevipalpis— 
appear to become rather a nuisance. 


340 Cc. F. M. SWYNNERTON. 


It is doubtful whether brevipalpis, a relatively easily detected and easily caught 
fly, would readily adapt itself to man. With regard to morsttans it may be said 
that the conditions favouring its dry season centres would have to be present as 
well as the villages. I only saw one such apparent coincidence here, but there is 
no reason why it should not be common. The size of the human population and 
its effect on the bush when really large are further factors to be considered. 


Of our fly areas here the “ Oblong” is uninhabited and the higher Mafusi and 
morsitans areas are not thickly inhabited. I believe it is very different on the actual 
Lusitu, but there brevipalpis would be the fly permanently present. 


I was myself bitten by morsiians far less than my natives, but this was presumably 
a matter of skin-colour and clothes ; Mabuzana, living near the Mtshanedzi south 
of Gogoyo’s, volunteered the information that tsetses were specially attracted by 
a black coat. I said “ How do you know that?” “ Because I have one!” 


Perhaps more surprising than the preferences shown in relation to man are those 
shown with regard to his domestic animals. I had three female native goats with 
me throughout the trip and in about the fifth week added a male; and until I 
obtained cattle, about a week later still, I used these goats continuously for bait. 
They proved most unsatisfactory, and although they were occasionally bitten, 
I bave little hesitation in saying that even man was much preferred to them both 
by morsitans and pallidipes and was not liked worse than they were by brevpalpis. 
Put ahead of the cattle they failed to draw fly where the cattle would draw many, 
A dog that was with me throughout the trip and still remains healthy was relatively 
httle bitten. Healthy-looking native dogs and goats were present throughout 
the fly-areas and were stated to be much bitten in the rains. I regularly fed my 
flies on the goats and once they bit they fed fully, but a relative reluctance to bite 
was distinctly seen, and this reluctance extended to a sheep on which I fed the flies 
that I finally brought home with me. The relative reluctance shown by Dr. J. W. S. 
Macfie’s flies to feed on guinea-pigs, dogs and cats is worth recalling, also the 
preference of Lloyd’s flies for monkeys as against goats. 


Experiments that I carried out on AstLIpAE showed that those flies possess 
graded preferences along the same lines as carnivorous vertebrate animals 
experimented on, so that there would be nothing extraordinary in the tsetses doing 
the same. Experiments on carnivorous animals, including lions, showed the 
following order of preference :—({1) Pig, wild and domestic ; (2) beef (nearly equal 
to pig); bushbuck and sable antelope were liked very nearly as much ; (3) goat, 
sheep, dog, man, blue duiker. These last were liked far less than (2). It seems 
probable from the evidence I obtained on this trip that the preferences of the 
tsetses I had to do with would follow much the same general lines. 


ASILIDAE often attack the highly nauseous Acraeine butterflies and tsetses attack 
man, but their preferences are very real nevertheless, in view of the fact that after 
a feed they do not wait till hungry enough for Acraea or man before feeding again 
should anything better turn up. Lloyd and others have shown, and my own 
observations confirm them, that when there is nothing better to turn up, the fly 
will at last “occur in such quantities ’—that is to man—“‘as to be a source of 
the greatest annoyance’ (Maugham). At Kanyezi’s vleis, with some game about 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 341 


but buffalos absent, we saw a good deal of fly and were often attacked by “ feeders ” 
(as distinguished from “ followers”). In the Masando vleis in 1900 with game 
temporarily very scarce and bufialos absent I was very greatly pestered. This 
time, in the same Masando vleis, with buffalos present, I found few flies, and at a 
series of vleis a few miles further east which the bufialos had just reached in numbers 
I found none, though the natives stated that they had been there in great quantities 
up till then. At first at these places I had not the cattle, and it was legitimate to 
suppose, as the natives did, and it was doubtless true of the males, that the tsetses 
were “away following the game.” But the female fly is a feeder, and apparently 
follows little, and the cattle quickly showed that flies were present, but that they 
were being kept too well fed to show themselves to less acceptable man. 

It is easy, understanding the working of the fly’s preferences, to see that a failure 
to find non-mammalian blood in an examination of its blood-contents is quite 
compatible with the possibility that it might live on birds and reptiles were mam- 
mals removed. 

It follows also from my observations on this expedition that the old idea that 
tsetses possess a preference for the bufialo may be perfectly correct, though it will 
show itself strongly only where, and while, buffalos are so abundant as to make 
the fly comparatively independent of less favoured food. The special enthusiasm 
I saw shown for an actual congener of the buffalo seems to be highly significant 
in this connection. 

Unfortunately there would seem to be no very practical method of utilizing the 
flies’ preferences except by using cattle to draw them out. With its favourites 
destroyed it will live well on the next best and so on. Arsenic-tolerant cattle, 
sufficiently saturated, might be worth experimenting with as bait. 


Drinking of Water by Tsetses. 

My flies frequently applied their proboscides to grass-stems put into the tube 
wet (to simulate the effect of dew), never to those inserted dry. My native 
informants on the Umvuazi agreed in asserting that they frequently saw tsetses in 
numbers “ drinking” on the wet sand of the river in very hot weather. I have 
watched Tabanus on the wet mud of pig wallows at the Amanzimhlope head-waters 
with its proboscis in definite contact with the moisture, as were those of non-biting 
Muscid flies of several species that were also present, and this habit of drinking 
is @ common one in hot weather on the part of both Diptera and butterflies. King- 
horn and Lloyd have both noticed tsetses at the edges of puddles ; Moiser obtained 
records of it from his natives, and saw his captive flies insert their proboscides into 
the wet soil of their bottles ; and Lloyd obtained the same result from moistened 
sponge and moistened blotting paper that I obtained from wet grass. Carpenter 
has gone further, for he has traced the presence of the liquid inside the fly. The 
point is not merely of importance in relation to the hot-weather needs of the fly ; 
it may also be important, it seems to me, in relation to the possibility of poisoning 
the highly localised male swarms of (. morsitans by spraying the grass they rest on. 

Lloyd’s positive results from slices of water-melon go to confirm Maugham’s obser- 
vations (under natural conditions) as to the sucking of vegetable juices. It is probably 
moisture rather than real nourishment that would thus be sought and obtained. 


342 C. F. M. SWYNNERTON. 


[X —DISTRIBUTORS OF THE FLY. 


Game not only helps (a) to feed the fly, and (6) to provide the trypanosome, 
but (c) it helps to distribute the fly, carrying it back each summer into the areas 
from which the fall of leaf had driven it. 


That the fly does definitely spread at each raimy season, over the latter type of 
area is Shown by native statements, by nagana outbreaks, and by my finding in the 
highly deciduous country a mudstained pupa-case of G. brevipalpis, one of our 
most shade-loving tsetses. That the fly wanders back into these areas partly 
without the aid of game is possible enough ; but without their aid I doubt if it 
would have re-occupied much country by the time the leaf fell again. The tsetse 
is a distinctly sedentary insect, and throughout reminded me much of a tick, 
excelling it mainly in flying a little distance to its prey instead of waiting to be 
brushed off by it. This, with the details of some of the outbreaks of trypanosomiasis, 
the fact that tsetses actually do follow animals for long distances, and the cogent 
indirect evidence I shall refer to on page 358, support the view that the game is. 
of the first importance to the redistribution of the fly. 


Accepting the hypothesis, the most important pomt to note is that the fly’s 
chief distributors will be by no means necessarily identical, in a given area, with 
its chief food animals. The individuals, pairs or herds of most of our antelopes 
have definite circumscribed haunts and grazing grounds that they keep to. In 
the case of bushbucks and duikers these are small, and parties of pigs also for long 
periods together rootle and depredate from particular lairs or lair-areas. Harte- 
beests also have relatively small beats, and herds of sable antelope and kudu have 
their permanent grazing grounds, larger than these others, though the kudu is 
sometimes a wanderer. Animals with this localised habit may be regarded as 
the fly’s most reliable food-supply in its permanent haunts, even though some 
other animal may temporarily become more important through sheer numbers. 


Elands, on the contrary, are great wanderers ; elephants cover much ground ; 
and the buffalo, if it is on the whole less of a wanderer individually, and in the matter 
of long purposeful “treks” than the eland, nevertheless covers, with its great 
herds, very large grazing grounds, is continually wandering back and forth between 
its various centres inside and out of the permanent fly-areas, and breaks up into 
parties and individuals that, to judge from the spoor, leave little ground in the 
general range unvisited in the rainy season. Also, as I shall show elsewhere, the 
buffalo more than any other animal has been increasing its numbers and range 
here at a great rate for many years past. 


I believe then that these animals, and the buffalo in particular, are mainly res- 
ponsible for the annual spread of the fly, and that so far as we are concerned, in 
and near Mossurise, our main grievance against the game is not so much that it 
feeds the fly (which would be fed and contaminated in any case by the pigs) but 
that it carries it far and wide in the rainy season and so brings it into contact with 
the cattle. 


The effect of the difference in the travelling habits of the male and female flies 
must of course be duly taken into account. 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 343 
X.—EFFECT OF THE RINDERPEST. 


The effect of the rinderpest is of practical importance in relation to the question 
of destroying the game to starve the fly. 

In reply to my question, addressed to many natives, as to whether the fly became 
less after the rinderpest, I received two answers, one from south of the Lusitu, 
the other from north. It may be stated at once that both buffalo and fly are said 
to be much more abundant, and to have been so before the rinderpest, in the Brachy- 
stegia area north of the Lusitu than in that to its south (both of them on the gneiss 
and carrying both morsitans and pallidipes), so that the difference in the replies 
may perhaps be genuine. 

The reply from the former area (from the countries of Muchamba, Usambai and 
Udombe) was that the rinderpest was followed for two seasons only by a definite 
reduction in the numbers of the fly. In the second of these seasons it was already 
more numerous, in the third season and subsequently it was as numerous as ever. 
The buffalos were still scarce, so that if this statement is correct and there were 
also any real connection between the rinderpest and the fly’s reduction, one 
explanation might be that the fly had previously become specially dependent on the 
buffalo, owing to the latter’s immense relative numbers, and took a season or two 
to adapt itself completely to the habits of the other larger mammals. Wart-hogs 
and baboons were stated to have been particularly abundant. Another explanation 
will be suggested below. 

The answer I received from south of the Lusitu, in the fly-areas both east and 
west of the Sitatonga Hills, was that the rinderpest was followed by no noticeable 
reduction of the fly at all. One man stated on the contrary that they then bothered 
the natives terribly. Another made the quite sound suggestion that perhaps with 
their other food destroyed they attacked men more, so that, though fewer, they 
seemed as many as ever. All insisted that, though certain abundant species of 
game were killed off, plenty of other food remained, in the form of bush-pigs, wart- 
hogs, baboons and part of the larger game. 

To the above evidence I am able to add a little of my own. In 1900 I passed 
with carriers through a portion of the morsitans fly-area. Hunting on 3lst May 
and 1st June in and near the large wood and series of adjoining vleis called the 
“ Mahloka ” or “ Masando ” near the Umvuazi we were beset by very great numbers 
of tsetses, which also kept with us for considerable distances—far heavier attacks 
and “followings ” than any I experienced in 1918. At that time buffalos were 
still exceedingly rare as the result of the rinderpest of four years before. I saw 
no trace whatever of them during that trip—a shooting trip—and local natives 
I have consulted have confirmed the fact of their absence. I have already mentioned 
the loss of a beast beyond the limits of the present fly in 1900, and the capture 
in the same year of a tsetse south of Spungabera, just inside the edge of the present 
permanent fly. 

On the other hand, my information from north of the Lusitu, if reliable (it was 
given to the members of my native expedition thither, not to myself), certainly 
seems to suggest that in some fly-areas an effect may have been produced. The 
real, dense, busily breeding centres of a morsitans belt may in the dry season at 


344 Cc. F. M. SWYNNERTON. 


any rate and under certain local conditions, be exceedingly localised. It is a fact 
also that the two animals that I have suggested to be the fly’s most important 
carriers are amongst the three that suffered worst from the rinderpest. It is possible, 
therefore, that the actual effect of the rinderpest in certain areas may have been 
not directly to starve the fly, but to confine it the year through to a far greater 
extent to its dry season centres by the destruction of its chief carriers. The narrow- 
ing in this way of the area tapped for food im the wet season might react also on the 
numbers of the fly. The suggestion is tentative and perhaps may not be borne 
out in the belts actually reduced, especially if man, a carrier of morsitans, was 
present in numbers, living or passing, about the dry season centres. 

The failure of the rinderpest to destroy the fly here to any appreciable extent 
tells decisively against the view that the rinderpest blood was in some way poisonous 
to the fly—for all my mformants agreed as to its severity everywhere, except in a 
portion of Makwiana’s highlands. The animals sometimes died “ whole herds 
together,’ and I myself can witness indirectly to the effect on the buffalos. 

Its failure tells also against the hope that we may exterminate the fly in the 
Mossurise district by destroying the bigger game only. 


XI.—FACTORS DETERMINING THE PRESENCE OF TSETSES. 


Shade and Undergrowth. 

It would appear from the present and other observations that the fly must be 
protected from the drying effect of continuous sunlight either by a sufficient supply 
of shade, or by a readily accessible moisture supply, or by both; and that this, 
with food, is its primary desideratum. 


Glossina brevipalpis, so far as I have seen (and I was with it for some weeks in 
all), relies very greatly indeed on shade and is rarely found away from fairly heavy 
shade. The requisite degree of shading is provided by wooding with leafy under- 
growth. This may be either primary forest (in which the undergrowth is sometimes 
such as to give sufficient shelter alone (P!. xui, fig. 2) or secondary bush in leaf 
with thickets and sapling clumps below (PI. xii, fig. 2). Of these types the primary 
forest (Pl. ix; Pl.xin, fig. 2) and, im many places and most seasons, heavy Brachy- 
stegia bush are the best capable of carrying this tsetse through the dry season. It 
is not at all dependent on the presence of vleis, and I have found it waiting in all 
the thickets at some distance from water of any kind and in hot weather in 
September with the ground baked. It is true that it was in greater numbers near 
certain little streams than in the Brachystegia thickets, but (as results from the 
same type of forest away from streams appeared to show conclusively) this was only 
because those streams were lined with fringing forest of primary type. 


So dependent does the fly appear to be on good shade that, except in the early 
morning, after sunset and on dull days, it will leave animals it is on as soon as they 
emerge from the shady bush into the sunlight. On dull and rainy days it will follow 
freely into the most open country and at high noon, so that the term “ crepuscular,” 
which has been applied to this fly, is not altogether justified by my observations. 
Thermometer readings taken at the same time in primary forest, primary forest with 
its undergrowth cleared, and, thirdly, in a sapling thicket in Uapaca wooding along- 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 345 


side, were identical, yet brevipalpis had deserted the cleared piece (v. Expt. 1, p. 373). 
This, with a failure to take brevipalpis in some high, dark secondary wooding on the 
Sitatonga base, excepting at the occasional thickets, suggests that extra shade 
may not be the only advantage gained from thickets. Protection from drying 
winds, additional protection from the eyes of enemies (such as the thickets very 
definitely afford) and some advantage to the pupae are the three that occur to me, 
but I am convinced that shade is the chief consideration. 

Glossina morsitans was found in a very different kind of country. It was very 
obviously far less dependent on good shade than G. brevipalpis, and I failed to find 
even stray individuals in the densest forest types—but the stray individuals taken 
were in any case very few. It occurs both on the granite-gneiss and the basalt 
of the lowlands, the latter usually with very poor bush indeed (Combretum, etc., 
Pl. x, fig. 2) and the former with savannah forest of a poorer type than that of the 
sedimentary area, though in each of these two it 1s Brachystegia. On the basalt and 
on most of the gneiss this fly occurred during my visits only as very rare individuals, 
and on the basalt and away from vleis only males were taken. The granite-gneiss, 
however, carries (as I have related) very numerous vleis, many of them with 
permanent water, and in spite of much painstaking search elsewhere, it was only at 
these vleis (and then only at some of them) that morsitans was found in great 
numbers together and its breeding going on with some vigour in the dry season. 

Here the male flies occurred in the usual bands, containing in some cases at least 
a few hundred flies at a time on the short or shortish grass and sedge. They were 
sometimes out in the sunlight amongst or beyond the scattered chidsgwati shrubs 
and stunted Parinariums that separate the open vleis from the Brachystegia bush 
surrounding them, sometimes a little way in the Brachystegia bush itself, never very 
far from the vleis. Some of these vleis with well populated fly-pockets consisted 
of mere glades in the bush, in which there was now no definitely exposed water and 
in some of which, had I not dug, I would hardly have expected water; but bigger 
vleis were part of the general series to which they belonged, and even in the glades 
experimental digging always showed moisture near the surface and actual water 
not many feet below. Some places would be moist on the surface, especially in 
the early mornings, when dew also was present in such glades generally. The 
flies were found in rather special association with a low, heavily-headed sedge 
(Futrena), with which I found the local natives had also learned to associate them. 
This remained green after the grasses generally were dry and, with one or two 
low-growing associates, covered considerable areas at the edges of certain vleis. 
Almost the only low-growing plant that was transpiring at the time of my visit, 


and doubtless cooler than the dried-up species, it possibly offered a greater certainty 
of the deposition of dew. 


The stray flies, of which I took fewer of morsitans than of pallidipes, appeared 
(as I shall indicate for pallidipes) to be finding their way to the vleis and rivers 
by the end of July, with leaf-fall already producing a visible effect in dry areas 
and at the actual vlei-edges, which latter, however, were also coming back into 
leaf. The natives described to me the increase of the fly in the hot weather, its 
spread in the early rains and its abundance then at places where, even with game 
temporarily absent, one now found few or none. In general it would be difficult 


546 C. F. M. SWYNNERTON. 


to find a more fitting description of this morsitans-pallidipes area than has been 
given by Dr. Shircore of the “ proclaimed area” of Lake Nyasa (Bull. Ent. Res. 
v, pp. 87 and 88) ; although, owing to the lateness of the season in 1918 and the faet 
that grass-fires had now barely begun, I am not in a position to judge of the complete- 
ness and duration of the stripping of the drier country of the stray fly of which 
I seemed to have seen the beginning. This type of wooding comes back into leaf 
again very readily after the weather has warmed up, and it possesses an early 
element in the mutsatsas. The return of the leaf does not await the rains and the 
country 1s sometimes already very shady when it is otherwise absolutely at its 
driest. The relative value of these two factors—shade and moisture—in relation 
to the fly’s fresh redistribution should be interesting to study.* 


I did not find that G. morsitans appeared to be at all dependent on the presence of 
undergrowth. The bush in which it occurred was for the most part devoid of such 
growth, though some of it was itself low. JI mayadd here that in consequence of the 
very late fires of 1918, and of the fact that I worked the normally late-burning 
areas last, the whole investigation was carried out in unburned country (except 
for a few odd patches). Nevertheless I obtaimed no evidence in favour of the view, 
but much against it, that any of the flies concerned will live in open grass-country 
devoid of bush. Even the attacks of morsitens on the basalt were always in or beside 
shade, were it only that of a large shrub or a semi-leafless tree, and when noted 
definitely resting, either in my experimental net or in the field, this fly was always 
in the shade—under a log, on the shady side of a trunk, etc. A replete resting 
morsttans female that I disturbed repeatedly always settled again on the shady 
side of trees. I have seen waiting male clusters furthest from bush in large vleis, 
but there was some reason to suppose that the individual flies did not stay with 
the cluster indefinitely. 


Glossina pallidipes is distinctly more catholic here in its general tastes than either 
of the other two. It occurs both in the country favoured by brevipalpis and avoided 
by morsitans and in that favoured by morsitans and avoided by brevipalpis, as 
well as in wooding of its own; nor do the stray individuals, at any rate, avoid the 
extremes. Males were taken on the sparsely shaded basalt (Pl. x, fig. 2) m dry 
windy weather and with the leaf falling freely, and occasional individuals of both 
sexes were taken in primary forest dominated by Khaya nyasica in full leaf (Pl. ix). 


I do not feel that I disentangled its habits sufficiently from those of G. morsitans 
on the granite-gneiss. Here, in the Brachystegia bush, it appeared to occur in every 
place in which we found morsitans (Pl. xiv, fig. 2; Pl. xv, fig1). It was in great numbers 
only where morsitans was also numerous—namely at particular vleis and glades ; 
it was very sparse elsewhere, but less so than morsitans, and both flies (and once 
brevipalpis too) were on us or the cattle together. When pallidipes puparia were 
taken at all, they were under the same log as those of morsitans. 


* Mrs. Lawrence, writing for her husband well on in November and sending me flies, 
speaks of the pleasant and cooling effects of the first light rains—‘‘ but” (she adds) 
‘“ the tsetses ! they are abundant.’ Tsetses had reappeared with the return of the shade, 
but the advent of the rains would seem to have led either to their freer breeding out or 
(as I think, by making more general the conditions under which they could exist in 
comfort or safety) to their freer dispersal from their breeding centres. 


THE TSETSE PROBLEM IN NORTH MOSSURISE, 347 


A fact that is possibly of interest is that in my first week spent at the Kanyezi 
morsitans centre (28th June—-6th July) I saw and secured numbers of morsitans 
but relatively few pallidipes, though a good sprinkling of this fly was present. Dur- 
ing my second stay there (27th July—lst August) pallidipes had apparently been 
heavily reinforced and was now taken (both males and females) in greater numbers 
than morsitans. It did not seem that emergences could account for this difference ; 
the puparia, living and empty, taken in large numbers on each visit, contained 
an extraordinarily small proportion of those of pallidipes—perhaps one per cent.— 
though the bush generally was well searched. On the other hand the adult flies 
of both sexes, which at my first arrival were scattered more through the country 
generally, had latterly, with only two or three exceptions (these being on the basalt 
where moisture would be harder to find), been taken only at streams. There was 
a difference now in the fact that the country was drier, and that leaf-fall had com- 
menced and was in places appreciable at the time of my second visit, so that the 
evidence suggested in a remarkable manner that pallidvpes, previously to a larger 
extent scattered sparsely over the face of the Brachystegia area, had now congregated 
at the morsitans centres. It is perhaps difficult to disentangle this conclusion 
with certaimty from the fact that the cattle always drew, and were now drawing, 
pallidipes in much larger numbers than natives working without them; but on 
this occasion the latter were themselves securing a larger proportion of pallidipes 
than previously. I should say that on the granite-gneiss the presence of pallidipes 
was in no way dependent on that of woody undergrowth. 


West of the Sitatongas, where morsilans does not appear to occur at all, stray 
individuals of pallidipes were taken in very open bush—and in every type of bush— 
but it was never taken even two or three together except in Brachystegia and the 
less tall of the “‘ dense secondary” types. It certainly occurs in small numbers 
in the dry season throughout the “ Oblong” and its northward extension to the 
Buzi. It is even possible that this great Brachystegia block may be its main per- 
manent habitat here, and it is notable that my cattle sustained a severe attack 
from pallidipes on 27th July in a limited piece of Brachystegia wooding just north 
of the Dysart Concession—a wood that had previously supplied a pallidipes or two 
daily to Dr. Lawrence’s grass-cutters just outside it. I also took a few pallidipes 
between this station and Gogoyo’s kraal at a series of small vleis in very dry sandy 
country carrying a type of Brachystegia bush equal only to the poorest on the 
granite-gneiss and already losing leaf somewhat heavily, and was informed by a 
native formerly resident that this fly was exceptionally bad here in the rains. Apart 
from these instances my only real and repeated successes with this species in the 
“ Oblong ” area were in and near the piece of coppice shown in PI. xvii, ‘fig. 1 and 
representing a type of wooding that, resultmmg from native cultivation, is more 
abundant north of the Mtshanedzi. Here (north of this river) on a few occasions 
up to half a dozen—even a dozen—together showed themselves in such types 
as are illustrated in Pl. xi (rather low thicket) and Pl. xvii, fig. 2 (coppice). 
Largely on account of this predilection, partly no doubt on account of the 
greater scarcity of brevipalpis and absence of morsitans, pallidipes west of the 
Sitatongas appeared to be specially associated with the Puizisi-Mtshanedzi area, which 
caiTies a very great deal of wooding of these types. The coppice or fern-entwined 
(737) ¢ 


348 C. F. M. SWYNNERTON. 


bush might be moderately open (as 1t was in Makwiana’s Makubvu through the 
trampling of elephants and buffalos). Very frequent—almost constant—con- 
stituents of it were the small trees or large shrubs, Brachylaena rhodesiana (ipahla), 
Markhamia lanata (mubfeya) and Vernonia podocoma (mudambasese), the latter 
bemg replaced near the Sitatongas and across them by another woody Vernonia. 
Bersama niassae (muyahawa), Vitex eyles (mukubvu) and Conopharyngia elegans 
were not infrequent, but a great mixture of other species, both of the savannah 
forest and the primary forest outskirts, were commonly present, and where the tree 
element overtopped the lower layer the wooding became fit for brevipalpis. 

Generally speaking, brevipalpis needs coppice with overwood, while pallidipes 
prefers coppice without it, but in the right types—such as rather poor Brachystegia 
—it can apparently dispense with yet lower growth. Clearing experiment No. | 
(below, p. 373) is of possible mterest in this connection. Here pallidipes attacked 
the cattle somewhat freely in tall primary wooding in which the undergrowth had 
been cleared, though before the clearing only brevipalpis had been present ; but it 
may have come direct to the cattle from the wooding (Pl. xvii, fig. 2) outside. 
It is curious that im all my special trials on the Buzi for brevipalpis in wooding 
of the types shown in Pl. xiu, fig. 1, I never took pallidipes either, though 
I did take it here in places with low growth. It may possibly best be described as 
a ‘‘low-wooding fly,” the low wooding varying from mere coppice and bush 
savannah to poor Brachysteqia, etc., though better wooding is to some considerable 
extent utilised. 


All vleis that I came across west of the Sitatongas were searched for male clusters 
of this fly and the neighbouring—and other—wooding for breeding places, but 
unsuccessfully. Though stated to be abundant at times in the rains it is generally 
by far the scarcer of the two flies where it occurs with brevipalpis. The latter 
often appears to be scarcer through its reluctance to show itself to man. <A point 
to be referred to also is the fact that pallidipes did not occur in all suitable wooding. 
Much would be passed through without encountering more than a stray fly, if any ; 
then suddenly a simultaneous attack by three, four, half a dozen or more might 
take place, as already described. Water (streams or vleis) was usually present 
not far away, but this part of the country generally is well watered, and sumilar 
water in similar woods more often failed to produce pallidipes. A small dry season 
population in a large area, and gregarious habits, probably sums the matter up 
west of the hills, but a study of my notes shows also that even stray pallidipes were 
relatively seldom taken anywhere at any great distance from vleis, glades or streams. 

Of Glossina austent I have in my possession a male fly and about sixty puparia 
all taken in two localities on the Mtshanedzi, near its junction with the Buzi. Fifty- 
four of the puparia, empty and (judging by their muddiness) dating from the late 
rains, were taken in the (roughly) primary-type forest that lines the Rupisi thermal 
stream, from under logs. The Rupisi at this point, immediately above its junction 
with the Mtshanedzi, was still hot enough to give me a pleasant warm bath, and 
its temperature doubtless influences that of the fringing forest. The latter is of 
appreciable width, and is here dominated by Khaya nyasica and such trees as 
Trichilia. It is, to be exact, secondary forest under invasion and still contains 
fine tamarinds (7. indicus), Kigelia pinnata and other secondary constituents, 


THE TSETSE PROBLEM IN NORTH MOSSURISE. (a 


but the primary and primary-like elements—particularly Khaya nyasica, but also— 
Trichilia sp., Lecaniodiscus fraxinifolia, musando (a splendid leguminous tree 
unidentified, sometimes erroneously called mukarati), Oxyanthus gerrardi, etce.— 
now so dominate as approximately to reproduce the conditions illustrated in Plate ix. 
The woody undergrowth is not quite dense, and there is a leaf carpet, not a live one. 
Great blocks of a crystalline quartzite appear above the surface, and the soil, 
evidently derived from this rock, is red:and finely sandy but compact. 
Many lianas are present, and it was under an inter-coiled mass of these that the 
main batch of puparia (30) was found. 

Shortly before reaching the Rupisi I had passed the ruined homestead of the late 
Mr. W. H. C. Coward, formerly a hermit in these wilds. His head-boy, aptly named 
Long One, had given me much information with regard to the tsetses hereabouts 
and had accompanied me on to the Rupisi to show me, he said, a forest in which 
small tsetses, in company with large, were extraordinarily abundant and trouble- 
some in the rains. It proved to be the fringing forest I have described. The small 
flies, he said, would follow one out of the forest, the big ones (doubtless brevipalpis) 
often turned back on reaching its edge after humming and settling about one im 
the forest. He described the attacks of the small flies as surreptitious. 

From four to four and a half miles on, near an outcrop of rock that perhaps 
represented the extreme southern end of the Sitatongas, and already beside the 
Buzi below the junction of the Mtshanedzi, we came on a great up-rooted tree 3} 
feet thick, lying head downwards in a donga, and beside it the rootlings of pigs. 
Under it at a point at which it was seven inches from the ground, were found 9 empty 
puparia of Glossina austeni, in sandy soil. The conditions were sandy, though 
somewhat less so than on the Rupisi. 

The general country between and near these spots was open, and the grass was 
more or less short with stretches of long grass. The chief savannah tree north of 
the Rupisi was perhaps Terminalia sericea ; a Pterocarpus (near angolensis) also 
occurred and in places much thicket of Bauhinia galpini and petersiana. Afterwards 
Bauhinia reticulata came into evidence and, more and more, the upoza (Combretum 
sp.), together with a general approximation to the conditions of the basalt suitable 
only for morsitans and pallidipes. Along the rivers the dense forest wooding in 
many places prevailed. 

There is much game in this corner of the country (Gowana’s). Buffalo, sable 
antelope, eland, Lichtenstein hartebeest and roan antelope are all present, also 
a few wart-hogs, an alleged abundance of pigs and baboons, monkeys (C. pygerythrus 
in the tree-savannah, C. albogularis in the thicker forest, as also much blue duiker), 
waterbuck, reedbuck, and plenty of bushbuck and cane-rats. Vleis (not seen by 
myself) are stated to be present, but not many. 

The one fly (austenc) taken by us bit the native who was giving my breeding flies 
a feed from a goat just outside the edge of the heavy Rupisi wooding, and possibly 
thereby exhibited a preference for man as against goat. 


Moisture and Rainfall. 
Morning mists, very wetting to the leaf, occur not infrequently im the dry season 
between the Sitatongas and the British border, especially in the great river valleys. 
(737) €2 


350 C. F. M. SWYNNERTON. 


In the bare glades and vleis of the Brachystegia bush of the granite-gneiss not only 
is there a continuous evaporation in the dry season from a considerable under- 
ground water supply, but the nightly cooling is such as to produce regular dews— 
as it does also, to a lesser extent, even in the drier open spaces in the woodland. 
If, then, the sucking of moisture from wetted grasses which appeared to take place 
in my tubes should also take place in nature, we should have a probable partial 
explanation of the attachment of morsitans (and in the same area pallidipes) to the 
vicinity of vleis and glades. In one instance a large fly-cluster was observed to 
travel quite a mile and a half in the course of a few days entirely along a connected 
or but slightly disconnected series of small glades in the bush. 


In view of such evidence as is alreadyavailable (page 341) I rather strongly suspect 
that tsetses will be found to partake fairly frequently of water in one form or another, 
especially the species that do not rely on heavy shade, in which dew is also less. 
It is conceivable that the slight occasional snacks of blood taken by male tsetses 
when travelling are as often for moisture as for food. 


At any rate the pomt with regard to water-drinking deserves investigation, 
seeing that the practical application of a discovery that tsetses regularly drink 
dew might lie in the possibility of poisoning at least the male flies at their dry 


season concentrations—as one of several contributory measures for the flies’ destruc- 
tion there. , 


Rainfall probably affects the fly here mainly in so far as it influences the nature 
of the bush, the period of leaf-fall and (for morsitans and pallidipes) the permanence 
of the moisture in the vleis, etc. I found brevipalpis and pallidipes both east and 
west of the Sitatongas and on the British border, though we have here, undoubtedly. 
three different rainfalls. The confinement of morsitans to the smaller rainfall area 
east of the Sitatongas will be referred to below. 


Up to 10th September (the end of the investigation), no matter how apparently 
suitable the wooding might be, I found no brevipalpis and only once a palladipes in 
the immediate neighbourhood of river banks west of the Sitatongas ; I refer to such 
rivers as the Buzi, not mere overshaded streams like the Inyamarimu. Yet on get- 
ting a bit back from the river, fly (mainly brevipalpis) was found generally and in 
quantity and thence up the hills to the limit of the suitable bush. There seemed 
to be a definite avoidance of the big rivers—possibly a matter of the greater winter 
cold in their vicinity. With austeni it may be different. 


Elevation. 


The effect of elevation was tested for brevipalpis and morsitans. Forty pupae 
of the latter fly were sent, twenty each, to two friends living in the mountains at 
about 3,400 (G. D. Otterson) and 3,900 feet (J. W. Scott), respectively. Each 
batch was placed in a little sand and leaf-mould (taken from the places I found 
them in) in a wide-mouthed bottle covered with gauze. Each bottle was placed 
under a slightly raised log in semi-shade, this reproducing the natural conditions, 


and the temperature was taken daily. I kept a control set in a similar bottle in 
the lowlands. 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 351 


The mountain sets were sent up late in June and more than a month of very cold 
weather supervened, including, it is stated, the coldest that we have had for some 
years. Without going into the details here it may be said that the tsetses of the low 
veld control showéd no advantage over those of the highland experiments. The 
percentage of failures was approximately the same, and the emergences both of the 
tsetses and of their parasites took place as well in the mountains as below. This 
is perhaps not surprising when one considers the elevation at which morsitans 
occurs in Mashonaland. 

I could only test brevipalpis observationally. On 27th-28th August, the colder 
weather being just over, I climbed the hills overlooking the Inyamadzi towards 
Spungabera up which the continuous Brachystegia bush extends to a considerable 
elevation, and I found brevipalpis present on the highest points at which bush 
conditions (except in certain inaccessible kloofs) were suitable for that fly—that 
is, so far as I could judge from the known elevation of the valley bottom, at an 
altitude of about 3,000 feet. It was steep and rugged here, but tracks of pigs and 
baboons were present. I understand that Jack took a pallidipes in August 1917, 
almost at the level of Spungabera, but the place (pomted out to me by 
Dr. Lawrence) suggests that it was a “carried” fly stranded there. In these 
observations on brevipalpis there was no break in the fly’s continuity, and from 
one to several flies attacked from the thickets right along from the Buzi to these 
high points near the border and back. 

I have already referred to my observation at Makwiana’s escarpment (fly at the 
bottom but none at the top), but this was simply a matter of shade versus none. 
The cattle sustained early morning attacks (before sunrise) from both brevipalpis 
and pallidipes with the thermometer at 47° F. and under. G. austeni was taken at 
an elevation of (probably) six or seven hundred feet. 


Shadeless Barriers. 

The isolation of patches of woodland suitable for the tsetse seemed of some 
importance in relation to G. brevipalpis in Brachystegia bush on the extreme edge 
of its area and where the intervening bush was already stripped of shade. 

Ascending at the end of August the slopes from the upper Buzi or Inyamadzi, 
or merely passing along near the bottom of the valley of the latter river with no 
alteration in elevation, one came out into completely leafless bush (Pterocarpus) 
or open grass, and | failed to find any fly at all in kloof strips or Brachystegia groves 
isolated in or by these shadeless stretches, although they were certainly too narrow 
on the Inyamadzi to act as a definite barrier to either sex. 

The obvious explanation seemed to lie in the refusal of brevipalpis to venture 
outside continuous shady bush except under certain conditions of light not common 
in the dry season, and then probably only if it can find a non-human carrier to 
take it. Unfortunately for this view and the practical suggestion arising from it, 
throughout the parts traversed of the low-lying crescent (Umzila’s guard-area) 
that descends the Mtshanedzi and swings back up the Puizisi, the smallest patches 
of dense forest with shrub growth below were very apt to shelter brevipalpis, few 
or many. The fact that the intervening bush was not yet leafless (in August) 
doubtless helped, and in places (as on the road east of the Chikambwe) this bush 


352 C. F. M. SWYNNERTON. 


itself carried some brevipalpis ; but in Mtobe’s area the intervening spaces were 
largely mere grass or bare Plerocarpus as on the Inyamadzi, yet the scattered 
darker thickets carried unusual numbers of the big fly. The clue is doubtless 
given in part by the fact that an actual detached primary-forest*clump, diminutive 
enough, near the Inyamadzi-Chiredza junction did produce a few brevipalpis and 
that the brevipalpis-producing clumps in Mtobe’s, etc., were of the shadier dense 
secondary or more or less primary types. The temperature conditions in continuous 
bush are more equable than in isolated patches, and the conditions for the existence 
of the fly may have been lost in such Brachystegia patches, though retained by quite 
diminutive clumps of the denser types. The Brachystegia bush on the Inyamadzi 
is rather poorer in any case in thickets and in fly than that across the Buzi, and 
it is right on the edge of the fly area with no passage to points beyond. Even 
if the explanation suggested is the correct one—and it must be noted that the 
observations it is based on were scanty—splitting up the shelter, as a substitute for 
complete clearing, on the margin of such an area would (by itself) exclude the fly only 
during a small portion of the year. 


Interesting also was the fact that kloofs ascending towards Spungabera from 
the infested Brachystegia wooding on the Buzi showed no brevipalyis appreciably 
above the general margin of that wooding, though much of the wooding in the 
kloofs was apparently still suitable for it, and I had taken this fly up to a greater 
elevation near the Inyamadzi. The explanation that this is due to the necessity 
for a compact breeding population (possibly in part applicable to the occurrence of 
morsttans centres in some vleis but not others) seems discounted by what I have 
just related of Mtobe’s country ; but this principle would doubtless show best 
on the margin of a fly area and if there were no passage of game to carry the flies 
to points outside. It was perhaps significant here that below the Spungabera 
forest a line of precipices divided the general fly-bush below from a wood of Brachy- 
stegia above in which there was much leaf and undergrowth but no fly, and that 
further along northwards the limit of the fly at that moment was not quite that of 
the Uapaca-Brachystegia bush itself but a little inside it at the foot of a precipitous 
slope up which the Gogoyo-Spungabera path goes, and on which the above bush 
terminates. The absence of fly from the Brachystegia bush above the precipice 
might be explained in any case on the lines that 1 have suggested for the isolated 
patches on the Inyamadzi, but the precipice very definitely and the steep slopes 
to some extent do seem to mark the limit here of the freer wandering of the pigs 
and other large mammals ; though pigs and baboons are said to come up in numbers 
in October and November to eat the ripening manzhanzhe (Uapaca kurkiana) fruits, 
and bufialos come up in the rains (especially from south and north of the more 
precipitous barrier) and, as the indirect evidence shows, bring up tsetses. I have 
been interested to hear from Mr. A. J. Orner, who has been along the path there 
recently in the rains, that even at that season, though he saw brevipalpis all the 
way from the Umtwadza to the foot of the steep ascent (mostly male queues in the 
path after sunset), after commencing to climb he saw not one more. In the rainy 
season there is much leafy wooding with thickets above the Brachystegia also, 
thus adding to the interest of an observation that is, however, incomplete through 
the absence of cattle bait to draw the fly out. 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 353 


Grass Fires, Grass Growth and Soil. 

The fires scorch the leaves and undoubtedly drive out the tsetses for the time 
being from the area burned, but on the granite-gneiss, on 27th July, I found 
morsitans had returned to a patch of ground that had been fired on 5th July. 
The grass was already 6-8 inches high even in the drier vleis and the trees on 
their edges, stripped by the fire, were already coming back into leaf. 

Of brevipalpis it must be said that of the two types of forest that it inhabits, one 
(primary forest and its extensions) never burns save along its edges, except in 
special circumstances which I have referred to already. 

With one possible exception, I should say that the tsetse pupa has no necessities 
such as are not found incidentally in the environment necessary to the adult fly. 
The nature of the surface soil is, I judge, a matter of nearly complete mdifference 
as a direct factor, for some slight accumulation of detritus is almost always present 
in such spots as receive the larvae, and I have found puparia on all soils. As an 
indirect factor it may be far more important, for it is quite certain that at most 
only an occasional pupa could survive the extraordinarily fierce fires of the heavily 
grassed dolerite outside the denser woods, when they do not take place too early. 


Food Supply. ° 

I have already referred to the réle of the big game in annually re-distributing 
the fly. I found no country in which largish mammals (pigs, bluebucks, etc.) were 
entirely absent, so that | am unable to say from my own observation that the fly 
can or cannot exist without them. That man must contribute much to the seasonal 
dispersals of G. morsitans I had abundant evidence, and I had plenty of evidence 
also that it does not avoid native kraals if trees are present. 


Two apparent Fly-barriers in Mossurise. 

(1). The Sitatonga Hills. These (except possibly at their extreme southern end) 
form the eastern boundary of the morsitans area. That this boundary is permanent 
there can be no doubt. It was the boundary when I visited the area in 1900 and it 
still is to-day. I failed to find the fly or its pupa west of it, and no morsitans has 
been amongst the numerous flies (brevipalpis and pallidipes) that have been collected 
for me durmg many months by Dr. Lawrence. Mr. Jack, who passed from the 
Gogoyo (Dysart) station to Mtobe’s, under the Sitatongas, in 1917, also took no 
morsitans. 

The Sitatonga Hills are a lofty and uninterrupted “ knife-edge ” passing right 
from the Lusitu to the Buzi. There is relatively little back and forth movement 
of game except at the southern end (where the buffalos have a passage), and it might 
appear therefore that the ridge acts as a barrier, either direct or (through the game) 
indirect. On the other hand, at Umtomana’s pass and about it the hills are very 
passable by game and have probably constantly been crossed by them formerly 
and are crossed to some extent now, and a main native path goes through here. 
My temperature experiment (above) suggests that the mere height of the hills will 
not constitute a direct barrier; and (a conclusive argument) the ridge evidently 
acts in no way as a barrier to the distribution of pallidipes and brevipalpis, which 
occur freely on both sides of it ; why then should it to morsitans ? 


304 


We must look for an explanation to a difference in the conditions. 


follows :— 


G. morsitans present. 

1. Smaller rainfall. 

2. Surface soil loose coarse sand on gneiss, 
compact on basalt. 

3. On gneiss impervious kaolin  sub- 
stratum bottoming numerous vleis, the 
latter fringed merely with the Brachystegia 
of the general wooding. 

4. Poor clean-stemmed savannah forest 
(Brachystegia) on gneiss; scattered bush 
savannah on basalt ; each type continuous 
over a great area. 


C. F. M. SWYNNERTON. 


They are as 


G. morsitans absent. 
1. Larger rainfall. 
2. Souls compact, much humus in denser 
wooding. 
3. Vleis fewer, mostly small and more 
often with long grass ; permanent streams 
numerous, largely fringed with big trees. 


4. Brachystegia savannah forest on 
sedimentary rock far finer and with, in 
general, much more undergrowth; on 
dolerite in widely scattered groves. Here 


the main wooding is (a) dense secondary 
types, also primary ; (b) highly deciduous 
tree savannah. 

5. Grass of tree savannah immense, 
fires fierce if late, milder in the denser 
wooding. 

6. Except in “‘ Oblong”’ less big game, 
but many pigs, etc. 


5. Grass mostly short, fires not severe. 


6. Much big game. 


2 and 6 may be dismissed. I found successfully emerged morsitans puparia on the 
compactest of soil on the basalt and the other two tsetses seem able to subsist here 
in numbers in the absence of much “big” game, as morsitans also did after the 
rinderpest. 


4 and 5. If (as is likely) it should be that only the tree savannah of the dolerite 
is really, as a whole, suited to morsitans, then its high deciduousness, with its 
fierce fires, should suffice to exclude this fly except as a possible summer visitor 
carried in on game or man. The scattered Brachystegia groves of the dolerite are 
utilised by pallidipes, and it is likely enough that it retires to these when the dense 
secondary wooding loses leaf towards the end of the dry season ; but to morsitans, 
unable to use the bulk of the bush, these Brachystegia groves would stand in the 
same relation as the outliers of the Brachystegia of the higher Buzi in which I failed 
to find brevipalpis do to the latter—only they are much farther away from the main 
block and separated from it by a high hill range and the two narrow and broken strips 
of dense, unsuitable wooding that fringe its eastern and western foot. To this extent 
the Sitatongas are a barrier, and doubtless also if they were more freely crossed by game 
morsitans would appear more frequently on their west side im summer. The apparent 
absence of morsitans from the generality of the continuous Brachystegia wooding 
of the “ Oblong” (neither Dr. Lawrence nor I have obtained any thence) seems 
to suggest that the difference in shade and undergrowth between this type of wood- 
land and the poorer type on the gneiss is sufficient to exclude this fly. 

To be able to assess the influence of the rainfall here one would require to have 
figures from many morsitans areas. It certainly seemed to me that morsitans, 
on the gneiss, was very dependent on vleis of a particular type for its dry-season 
breeding. Vleis quite of this type are rare, if not absent, on the west side of the 
hills, but it is again difficult, without an acquaintance with other morsitans areas, 
to know if these vleis do really represent a vital condition. It may be this factor, 
rather than that of the wooding, which causes the demarcation, and at present 
that is rather strongly my opinion. 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 39D 


In general it does not seem likely that the country west of the Sitatongas, at any 
rate north of a line drawn through the middle of the “ Oblong” and parallel 
to its greatest length, is in any danger of being invaded by morsitans. Occasional 
flies might be carried over the hills by Kafirs or game, and the latter would probably 
happen more frequently were the kraals on the eastern foot of the hills to be removed. 

(2). Lhe mountains of and near the British border. These constitute the western 
and northern boundaries of the pallidipes-brevipalpis areas, and for a clear idea of 
them the map should be consulted. Generally speaking they form more or less 
of an escarpment, abutting on to the fly country on these two sides. 

The great rise in elevation seems at first sight an adequate reason for the existence 
of the fly boundary, which, it should be repeated, appears always to have stood 
approximately where it is now, except when pushed east by Kafir settlement. No 
doubt the British hills north of the Lusitu (6,000 feet) and the Chimanimani (much 
higher) would act in this way as a direct barrier, but it is more than doubtful 
if, from the Lusitu southwards, the elevation (with maximum heights of a little over 
4,000 feet) is sufficient to act as a direct deterrent to the fly. On 
the contrary, a glance at the map shows clearly that it is the great continuous 
pieces of wooding of the right types that really delimit the fly. Where these 
ascend mountains, the fly ascends equally ; where they stop short in low-lying 
valleys (Inyamadzi, Chiredza, Buzi, Lusitu), the fly stops short too. It happens that 
these great pieces of wooding cross the British border for a short distance only 
where they touch it. With regard to the wet season fly, which is capable of being 
established temporarily by game in occasional pieces of wooding such as exist 
abundantly enough even on the British side of the border, another factor must be 
mentioned—the white settlers on the border and such native kraals as are present. 
Our experience shows that these do act quite considerably as a deterrent to the passage 
of the wandering big game, and that where a white man shoots much or native 
kraals are numerous a useful “ guard” against the fly is thereby established. 

In general, however, the summing-up of old Inyabangwa, who long ago herded 
cattle for the Zulus on the edge of the escarpment, fits the case. “ They are barred, 
sir,” he exclaimed, “barred (zi-vimbekile, ‘nkosi, zi-vimbekile). They fear the 
open country (izimbaya) of the mountains and they fear its cold winds.” 


XII.—Nores on Hasirts. 
Waiting Habits. 

I have referred to the tick-like habits of tsetse. I came across no evidence that 
the flies wander to any extent in search of their prey, though a good deal of local 
activity is shown by male brevipalpis after sunset, and females then attack from a 
greater distance. That some of the stray flies found in the country generally are 
earried, I had evidence in the sudden appearance of my marked flies in native 
villages immediately after parties of reed-cutters had passed through them from my 
vleis. 

G. brevipalpis may be found in the day-time scattered through all the little 
thickets in the bush it frequents, two or three or more to each. Just at sunset the 
males emerge from the thickets in the neighbourhood of game and other paths and 
(where plentiful) distribute themselves at short intervals along them, sometimes 


356 C. F. M. SWYNNERTON. ¥ 


for a considerable distance. As one walks each male moves on in front of one for 
at most a few feet, evidently scanning the walker for any females that may be with 
him, then falls behind. At one time when a domestic matter called me home from 
the Buzi, my police boy, left in charge of the work, captured no less than 88 of these 
flies, every one a male, on three successive evenings without the aid of cattle along 
the same stretch of path. The path here passed alternately through bush suitable 
to brevipalpis and more open spaces. On emerging into the latter one ceased to 
see flies; on entering bush further on one found them in the path. This was a little 
curious seeing that it was after sunset and rather suggested that they had not come 
far. 

The males of G. morsitans, though occasionally seen like this on game paths, 
were found mostly crowded together in the short grass beside the path—some of 
them on the path—and were also (as events each time fully proved) watching for 
females or carriers to take them to the females. G. morsitans usually waited in 
a crowd, brevipalpis always in a queue. 


Travelling Habits. 

G. morsitans utilises man as a carrier far more freely than the big fly does, and 
as these followers occasionally bite, they would give an unobservant person the 
impression of persistent attacks by large numbers of tsetse. It is, actually, nothing 
of the kind, but, in essence, a ride in search of females. This is most clearly seen 
when the flies are few and on the back of the person in front. I have watched 
male flies, each with its station on a particular spot on the back of my guide, travel 
thus for miles. They would keep taking short flights off all the time and at once 
return to their previous station. Whenever I saw the object of the flight it was 
another fly, and the fact that the flights were in search of females was sometimes 
proved by a witnessed pairing. 

That these rides on man would be useful even where the females were not feeding 
on man was supported by the many instances | saw in which flies came out to us 
and turned back, evidentally rejecting us. (These observations were on brevipalpis.) 

Where I have walked slowly the tsetses (morsitans) have some times contented 
themselves with moving along in the grass beside me. More often they—or some 
of them—attached themselves to me when I was moving and each time I stopped 
dropped to the grass again till a movement showed it was time to start. There 
can be little doubt that when on the grass they watched for females coming to me, 
for such females were at once assailed. The low position adopted by the tsetses 
during these waits was doubtless of considerable use in showmg them the females 
against the sky, and in helping them to detect and keep in touch with moving 
objects generally. Fhes that had flown off a man’s back to accost another fly 
always returned flying very low and rose to his back. The principle is that of 
the nightjar or of the man who stoops to watch crepuscular insects or birds. These 
clusters of male flies varied greatly from day to day in numbers and position, 
reductions definitely resulting both from the depredations of birds and from the 
flies attaching themselves to passers by. With buffalos present the clusters 
practically became non-existent. They always tended to form again later, but 
were not always composed of the same individuals, for experiments with marked 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 357 


flies showed that these reappeared in other clusters than those in which they were 
marked ; and the immense preponderance of unmarked flies in clusters that I had 
taken pains to mark pretty fully suggested that they were being kept up not merely 
by arrivals brought from elsewhere and from males that had probably been resting, 
but by new emergences in the neighbourhood, such as were actually noted in searching 
for pupae. In each case active breeding centres and clusters were never found 
otherwise than in more or less close contiguity. 

There was a certain amount of day-to-day movement on the part of some of 
these clusters as a whole, but it was gradual. The one nearest my Kanyezi camp 
Was very stationary, its movements being mere fluctuations, and after a three 
weeks absence I was able to find both this and the other clusters almost where I 
had left them and to retake in them a few of my marked flies. 


Males of brevipalms waiting on game paths were frequently tempted to attach 
themselves to the cattle and would then often ride for quite a distance. Sometimes 
they fell behind them, as they fell behind ourselves, probably because they found 
that they carried no females. 


I measured one of these rides by five male brevipalpis in dull weather, first marking 
each of them with a dab of white paint. The flies had come to us just before and 
were following, not feeding. The greatest distance travelled was 54 miles by the 
one fly that stayed to the end. All five flies were still present at 4 miles and 700 
yards. Every kind of country was traversed, brevipalpis bush, simple coppice 
and open grass. 


The flies deserted us on our getting back into their native bush. Elsewhere 
where brevipalpis had followed us into the open in the daytime it waited before 
leaving us until it had reached suitable bush, and an individual that once followed 
far to a camp in the open stayed about for hours (in the evening) before disappearing. 
All this was interesting in relation to the explanation of outbreaks of nagana in 
which it appeared that the fly had crossed much open country to bush—patches 
in the grazing grounds or near a kraal. 


I tested similarly the d'stance to which morsitans will follow man. Twice I 
obtained rather more than six miles, and many times various less, but still long, 
distances. On one occasion a journey of over six miles was done in three instalments 
by a marked fly on three successive days. The riders were almost always inter- 
rupted by reaching, sooner or later, another clump of male flies into which the male 
riders then disappeared. Otherwise, and with more time to experiment, the rides 
might well have been much longer, and Bevan (I think) has recorded an observation 
of Meikle’s in which tsetses were still on his cattle 25 miles outside the fly-belt. 
A common position was on the load of a native carrier. 


In the foregoing experiments and observations the flies that really travelled- 
were invariably the males (marked with a different colour), and it was these only 
in the case of morsitans and pallidipes that had up to the time of my departure 
put i an appearance at villages a few miles outside the breeding centre in which 
they were both captured and released. Hungry flies, including, so far as I saw, 
all females, came to feed, not follow, and the longest distance travelled in an 
experiment in which I released a considerable number of hungry marked brevipalpis 


358 C. F. M. SWYNNERTON. 


and a few pallidipes behind the cattle was 820 paces quickly covered. The other 
flies had already dropped off replete before that, in spite of the kicking and running 
on the part of the cattle. 


The question of the females travelling, even on animals, is an important one 
in relation to the question of possible control by barrier clearings, and is worthy 
of more extended special experimentation if long distances have not yet been 
recorded for it.* In Simpson’s experiments with marked male and female flies 
on the Yapi-Tamale road, in the Gold Coast, only one recaptured fly was found 
more than two miles from the point of release, and all were retaken near the road, 
on which there was considerable trafic. Unluckily im stating the result he does 
not separate the sexes or state the time that elapsed between release and recapture. 
A female could readily cover two miles on successive journeys on man and animals, 
especially if much disturbed in her feeding. In Lamborn’s experiments only males 
were retaken. 


It is obvious that a fly-belt can only extend definitely, whether its annual extension 
into highly deciduous wooding is concerned, or a permanent extension into country 
previously uninvaded, at the same rate as the females. This, if I am right in my 
present belief that the females do not travel appreciably on their own initiative 
or to any distance on animals, suggests that the extension will be slow and compact 
and indicates a possible explanation for the alleged fact that the cause for the 
division between fly and non-fly country is sometimes not to be found in any differ- 
ence in the conditions of wooding. 


Living in a district that is right up against the fly I have come across no evidence 
at all that either sex travels appreciably on its own initiative. There is on the 
other hand, a very great deal of evidence to connect our local outbreaks of nagana 
with the movements of big game. I do not think that a fly area will extend seriously 
in the absence of game. 


Migration and Homing. 


The fact that the flies are found still in their centres in great numbers when food is 
scarce, and even under such conditions of hunger as have been described by Maugham 
and as, it is just possible, I saw myself in 1900,} tells against the theory of unassisted 
outward migration, though the predilection of the “following” males for joming 
other fly-clusters they may meet on the road amounts to unassisted migration of a kind. 
It is also quite certain that in South Melsetter, with the fly right on our border for 
20 years past and numerous herds of cattle grazing in and near Brachystegia bush 
at relatively low elevations, we should have met with much indirect evidence of 
migration were such to take place. 


* Against the obvious suggestion that the barrier could be narrow must be*placed 
the possibility, to be referred to below, that females as well as males do not dismount 
readily in uncongenial country once they have allowed themselves to be carried well {into 
it. This point needs testing also. 


+1 see from my journal of that time that the tsetses “‘swarmed andj were very 
irritating.’’ Game was not quite absent, but, with the exception of duikers, was very 
scarce, having left this unburned country for the green patches on the basalt that had 
been burnt earlier. 


THE TSETSE PROBLEM IN NORTH MOSSURISE., 359 


That homing occurs, within limits, is to me certain. On 29th June as soon as 
I entered a male crowd the flies as usual attached themselves to me and kept pace 
with me, buzzing about my feet. On my leaving the vlei and entering the Brachy- 
stegia their numbers fell off rapidly, only a few following me 50 yards in. I[ stood 
still finally when all had left, and after a few minutes returned to the cluster to see 
if I should pick up the flies along the path, where they had left me. I saw not one 
fly till I had got to the cluster, in which they seemed as numerous as before. It 
seemed probable that on leaving me they had flown straight back to the cluster. 
Natives arriving with loads shortly afterwards passed through the cluster and a 
number of the flies accompanied them right through to my camp—a further 150 
yards—possibly a matter of preference. They stayed for a short time, occasionally 
biting, then all disappeared. 

Similarly whenever we emerged from a cluster, followers would just at first often 
be numerous, but for the most part would fall off within a very few hundred yards, 
and on the analogy of the above observation and from certain takings of marked 
flies, it seemed likely that these short-distance flies had flown back. The numbers 
of the remainder lessened rapidly with the distance travelled and relatively few 
followed for some miles. Would these long-distance flies also find their way back 
unaided ? Would they necessarily proceed further than the first useful vlei? And if 
deposited in congenial surroundings, would they necessarily return at all? The 
experiments known to me on the subject of long-distance homing do not (as described) 
seem to settle conclusively the question as to whether it was unaided. Simpson’s 
were on highways of traffic. His recaptures were many and confined to the vicinity 
of these highways—road and river. Lamborn’s recaptures were few and his general 
distances longer. From my cattle experience I judge it to be likely that fly will go 
farther on gamethanon man. They also, in my observations, tended to travel farther 
through unattractive country, once there, than through country offering temptations 
to dismount. This suggests further the need for a closer description of the country 
searched on the wrong side of the point of release. The question is of some possible 
practical importance, and it was, therefore, a matter of regret to me that lack of 
time, with the fact that on first leaving the Kanyezi centre I did not know if I could 
return to it, prevented me from carrying out long-distance experiments of my own. 


My incidental evidence suggested that homing does not always take place at once. 
The fly that I have described as travelling six miles in three stages reattached itself 
each time at the spot at which it had left one or other of us the day before, and 
other isolated instances of the same thing occurred. Of my marked flies that were 
carried to native villages a few miles off two were reported as staying about for from 
two to four days before disappearing—when they may merely have moved farther 
on on natives or have been killed. Further evidence was obtained in investigating 
a severe and sudden outbreak of nagana in a large herd in British territory, very 
many miles from fly (permanent or otherwise), which appears to have been caused 
by male tsetses carried by game. It had taken place at the commencement of the 
dry season. The native in charge and the herd boy, interviewed separately, each 
picked out tsetses (from amongst my specimens) as having been on the cattle before 
and during the outbreak and stated that they had appeared suddenly and stayed three 
weeks or rather more. They disappeared “ when the leaves fell ’’—a spontaneous 


360 C. F. M. SWYNNERTON. 


piece of evidence which, coming from high-veld kaffirs previously unacquainted | 
with tsetse, seemed rather convincing. Whether they then “ homed” or died 
it is impossible to say. Old natives who herded cattle in the earlier days of Umzila’s 
experiment had previously given me similar instances of long stays by the fly. Of 
course there was an attraction, but the seasonal spread of the fly into other deciduous 
areas (v. Sections ix and xvii) will itself, if fully confirmed, constitute strong evidence 
against the universality of homing, as the well known recent spread of many fly 
areas already does against its completeness. 


Hours of Activity. 


G. brevipalpis, as I found, attacks in the shade of the bush at any hour, but does 
not fly appreciably outside its own thicket to do so except in the early morning, 
after sunset and on dull or rainy days. Having attacked, even in sunny weather, 
it completes its object of feeding or riding, unless its victim emerges into the open. 
In this event it usually leaves. In a special experiment that I carried out with the 
aid of a lantern occasionally lit and turned on I found that both this fly and pallidipes 
attacked right up to dark. Then every fly suddenly disappeared. On moonlight 
nights they continue their attacks, and all three flies come to light, including fire- 
light, and, once there, bite'freely. G. morsitans I found, as have other observers, to 
be most active in the rather warmer hours and least on cold or dewy mornings, but 
it too attacked up to dark. It is probably dependent on shade to retire to occasion- 
ally when blood and water are not available, for I captured several male flies that 
had been resting under logs in the wooding adjoining vlei edges frequented by male 
clusters. The frayed wings of some showed that they had not just emerged. 


The Bite, and the Attack; Methods of Marking. 

Nothing struck me more than the frequency with which a fly would steal a feed 
from the back or (especially) leg of a kafir walking in front of me and fly off replete 
before the man noticed it. Very frequently the first intimation of the fly’s presence 
comes with the final withdrawal of the proboscis ; sometimes even that is not 
noticed. At other times a bite is noticed instantly and felt acutely. The natives 
who converse on the phenomenon hold that a bite over a bone hurts and in a fleshy part 
does not, but it 1s obviously a matter of local presence or absence of the specific 
sensory nerve-terminations. I have myself had both painful and painless bites 
quite close together on my forearm. 

The practical bearing of the fact is in relation to the criticism (based on the success 
of observers’ pet animals) that flies will not make good their feeds on baboons, étc., 
and that such animals need not therefore be seriously considered in relation to the 
efiect of game destruction. A second fact to be remembered here is that captive 
animals have abundant leisure for fly-catching. A wild baboon is often absorbed in 
raiding. hastily cramming his pouches, quarrelling, keeping up with the troop, or 
participating in shouting contests. Especially where tsetses are present all the time, 
he can hardly attend to them except incidentally. The black man in these circum- 
stances brushes the fly he notices away—commonly, if walking, by a wrist move- 
ment that brings the butt of his assegai sharply to the point on his back that was 
bitten—and the fly at once returns ; he makes an efiort to capture it chiefly when 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 361 


he is idle. When watching tsetses in my net experiments I relatively seldom 
noticed the approach of any that attacked, though their attentions were sufficiently 
annoying to make me wish to do so. A male in search of females approaches boldly 
enough and is not silent, but the Geng of flies intent only on feeding is often 
remarkably surreptitious. 


I was never bitten through my clothing by G. morsitans, but very frequently 
indeed by hungry G. brevipalprs, both through thin shirt and vest and strong khaki 
trousers, though I occasionally watched flies probe for a time as though unable 
to penetrate. The scrotum (when one squatted) was almost always the point 
of attack, and this accorded well with this fly’s habit in relation to cattle. It 
always attached itself to their bellies, more or less in the median line, or between 
the fore or hind pair of limbs. The kicking of its belly by a beast in brevipalpis 
country almost invariably indicates that it has been attacked by that fly. The 
position chosen has probably as much to do with considerations of shade as with 
the thinness there of the skin, and some very hungry brevipalpis released by me 
behind running cattle fed at the first points reached, including the flanks, whence 
one was seen to be licked offi—suggesting a further reason for the selection of the 
belly by a fly that is less active in avoiding such reprisals than is G. morsitans. The 
smaller tsetses attacked more on the flanks and forequarters and lower down. 

I carried out an experiment in trapping tsetse by fastening skins smeared with 
bird-lime on the goats and under the animals’ bellies. The fly-catching properties 
of the lime, made from Loranthus berries, were excellent, but it speedily became 
so masked with leaves and grass blades as to be useless. 


I have referred already to the low poise and sidelong, running, almost Olfersia- 
like movements of the fly in evading reprisals. This is well seen when the natives 
try to catch it, as they frequently do, by “treading on its toes”’ with the edge 
of an assegal or knife. -Occasionally a leg is damaged in the process and this, with 
the attacks of birds, makes unreliable the method of marking by snipping off a 
portion of a leg that, was adopted at first. I later used paint from water-colour 
tubes, placing with « small brush a dab of white or colour on the back of the thorax 
and varying its position and size. This proved excellent. It enabled me in some 
cases to recognise individuals again and again, and the conspicuousness of the 
mark brought me reports from the natives as to the spots at which such flies 
reappeared. It doubtless also carried with it the disadvantage of laying the flies 
open to detection by birds—a consideration that is of less importance in relation 
to the males, which, in any case, trust more to activity and less to concealment 
than the other sex. 


XIII.—PROPORTION OF THE SEXES. 


The following statements represent the result of day-to-day observation and 
analysis in the field—the only reliable method. 


(1). Wherever we had to deal with male crowds or queues awaiting females or, 
in the case of brevipalpis, the bush immediately bordering on the path on which 


the male flies would line up at sunset, we took, as might be expected, practically 
nothing but males, 


362 Cc. F. M. SWYNNERTON. 


(2). Elsewhere, using cattle bait, we found usually either an approximate equality 
of the sexes or a preponderance of females. 


(3). Many more females proportionally came to cattle than to man and goats. 


The cattle were with me only durmg my last four days in the morsitans area 
(three at Kanyezi centre), but the proportion of females rose abruptly on their 
arrival. It amounted now to from 35 to 40 per cent. in spite of the fact that 
the cattle were used very largely at a strong male cluster im order to obtain 
the material for certain experiments. Previously 7 per cent. had constituted a 
good day, the only day on which I had surpassed this figure being one (1st July) 
on which the already weakened male cluster at and near which we were catching 
had temporarily been obliterated by the passing through it of a buffalo. The 
percentage of females for that day only was 30, and it was also the one day on 
my first visit on which pallidipes (taken one here, one there) predominated, but 
the total was in any case diminutive, not exceeding my daily catch minus morsitans 
males, 


Bagshawe’s result (more male palpalis where crocodiles were present, more 
females where not) and McConnell’s (more palpalis females on the small tributaries 
—where food is doubtless scarcer—more males on the main Nile) appear to be 
completely explained by observation No. 3, above. Bequaert’s observation (more 
males m uninhabited regions, where non-human food would be more abundant) 
would appear to be the same, had not Graham (quoted by Bagshawe) noted that, in 
Ashanti, palpalis prefers human blood to that of cattle. But both Graham and 
Zupitza (also quoted by Bagshawe) observed that when flies bite readily males 
are not in excess, 


In spite of a heavy preponderance of males in my total takings—the inevitable 
result of following paths and of special attention to all male crowds or queues 
with a view to obtaining material for my marking and other experiments—I came 
across no evidence at all which, properly analysed on the spot, suggested an actual 
preponderance of the male sex. Rather the reverse; and it was likely, both m 
view of the systematic attack by birds which I found taking place on a conspicuous 
male crowd and from a consideration of the breeding habits (which are such as to 
allow for much male wastage), that, at a given moment and apart even from the 
freer dispersal of the males, the females are more numerous than the latter, though 
they will show themselves in their true proportions only when hungry. 


Dr. Lawrence’s numerous small sendings of flies, lasting over many months, 
showed (in pallidipes) a remarkable equality of the sexes even in the individual 
sendings. Here we have a locality in which game generally is rather scarce and 
even wild pigs, it is stated, unusually so ; also no male fly crowds, in so far as I 
have seen or heard, but mere individual attacks by single flies on natives passing 
or working. In the stray individual flies of pallidipes and morsitans taken by my- 
self over a far greater variety of country both east and west of the Sitatongas, 
some of it with much game, the females amount approximately to one-third, but 
a few stray flies taken on cattle (on Ist Aug.—females over 50 per cent.) are included 
here. 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 363 


Attacks by several pallidipes together (except at the waiting male crowds of 
the granite-gneiss) were very different. Here the tendency to equality was great, 
and on at least one occasion, to judge by the captures, only females were present. 
But we had no such attacks except on the cattle. 

The above results from pallidipes. may ‘usefully be contrasted with those obtained 
from brevipalpis—a fly which is very loth to feed on man; and it must be remembered 
again that the female sex comes primarily to feed. In Dr. Lawrence’s sendings 
(mainly flies taken on natives) practically all the brevipalpis are males. Except 
in the pig observation (p. 337) this was my own result also, till I used cattle bait. 

My results would seem to place Lloyd’s suggestion outside the category of theory 
and further to show clearly that the apparent scarcity in the presence of game 
of the females of the three flies chiefly investigated is probablv due to a relative 
dislike of man. 

The fact that morsitans and pallidvpes might be found about native villages 
where the shade was suitable, was, as I shall show, the natural result of villages 
being centres of arrival from the surrounding country, together with the fact that 
these flies do attack and (in particular) follow man. It does not invalidate their 
probable preference for game. 


XIV.—OBSERVATIONS ON BREEDING HABITS. 


Glossina morsitans. 

Wherever I went I searched diligently for pupae with all the natives at my 
disposal, including often a number of local natives incited by the offer of rewards. 
Special halts were made at likely or typical spots or at mere fallen logs, and the 
normal halts were fully utilised. My general lack of success was remarkable, and 
the two or three solitary, evacuated, muddy, wet season pupae found corresponded 
with the sparsely scattered character of the actual fly in most of the morsitans— 
pallidipes area. Only one small find of puparia (evacuated) was made on the basalt 
in spite of a week’s exhaustive search in the country round my camp, both in 
river bush and bush savannah, by a large and quite keen gang. 

At some of the better vlei-series (by no means at all) the indications were very 
different, and at one of these (shown on the map as Kanyezi’s) I stayed at the end 
of June and again at the end of July, for about a week each time. Burffalos had been 
present here in very great numbers in the late rains, but had left two months before 
my first arrival. During my stay a small herd of Lichtenstein hartebeests, small 
droves of wart-hogs, and latterly a solitary bufialo bull were about the only large 
mammals present, except an occasional duiker, a leopard and three passing lions. 
The bufialos had left some heavily trodden-out game paths, now used mainly by 
ourselves and once by the solitary bull. Thorough searches for puparia were made 
both near the game-paths and away from them and near the vleis and away, and I 
took many hundreds of full and evacuated puparia in all from more than a hundred 
breeding places. The puparia were not, like Lloyd’s, necessarily connected with 
the game paths, though it was not easy to get very far away from these ; but the 
muddy ones were more or less confined to areas well grazed by the buffalos, and all 
were confined to the wooding in the immediate vicinity of open vleis and glades, 
(737 D 


364 Cc. F. M. SWYNNERTON. 


to the borders of which and the old game paths traversing them the male flies were 
obviously betaking themselves on emerging. The soil was sandy, the formation 
was granite-gneiss, and the wooding was the Brachysteqgia-dominated savannah 
forest of that formation. 


The females when not feeding certainly spent their time in hiding away—from 
the males (as Lamborn has suggested) and, I am certain myself, more especially 
from birds and the sun. The indirect evidence showed clearly that it was mostly 
when thus hidmg—in a spot which suited their own convenience but by no means 
necessarily that of the pupa (cf. also Lloyd) —that they dropped their young, sometimes 
(as my indirect evidence again most strongly suggested) from several feet from the 
ground. Most of my puparia were found under prostrate trunks and branches, 
raised little or much from the ground, but batches were also found in the angles of 
root-buttresses, under mere leaning trees, under fallen palm leaves, and below and 
between the outleaning dry leaf stems at the bases of palms (Hyphaene ventricosa), 
in holes in trees and in holes made by animals in banks, and where the hoof-marks of 
passing animals had broken the slight crust of a sandy stream-bed immediately 
under light overhanging grasses. I failed, in spite of much search, to find tsetse 
pupae in the leaf-sheaths of palms, though I took Lepidopterous and often other 
Dipterous pupae there. The situation was usually dry, but some of my sand-stream 
pupae were in a moist situation, and I took a pupa-case that had emerged success- 
fully under a fallen Hugenia log in very damp ground on the edge of a vlei. 


I also carried out an experiment, lasting many weeks, with the pupae taken. 
Some were exposed to conditions as normal as I could make them, some were in 
soil exposed to day-long sunlight and others were in ground kept always moist and 
either warm or cold. It was wonderful how well those exposed to the extremes 
(perhaps less extreme than the conditions used by Roubaud) emerged, but less so 
when one remembers the continuous state of the ground in such a wet season as 
1917-18 or so dry a one as 1912-13. Pupae placed in a calabash of sand exposed 
continuously to the sun failed, but in travelling they worked right down under the 
sand in any case and many of them showed definite crushing by it. It 1s worth 
noting that in the moister situations (in the vleis) practically no trees grow except 
en the ant-heaps, consequently logs and other dark hiding places are relatively 
seldom found there. Otherwise it is probable that pupae would have been taken 
there more frequently. 


One other condition should be mentioned. The vast majority of the pupae were 
in the nearest suitable hiding place to a spot in which big animals had been lying 
down—buffalos (mainly), wart-hogs, hartebeests, etc. This seemed to accord with 
the fact that we ourselves were attacked by feeding flies (as opposed to males 
utilising us as vehicles) mainly when we were stationary and especially when we 
were squatting down searching for pupae. Again, of two sand-streams, otherwise 
not dissimilar, one that was used as an occasional game path yielded a number of 
pupae; the other, used only by small carnivora, yielded only one pupa—in a highly 
placed hole where a duiker had lain on the edge of the bank just above. It was 
frequent for the puparia to be near ant-heaps, but this was the result of the fact 
that the buffalos had chosen the sheltered sides of ant-heaps to lie on during the rains. 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 365 


Provided that the hiding-place itself afforded shade to the mother, overhead shade 
(as an analysis showed) was a matter of complete indifference, and the ground in 
which puparia lay was commonly reached by the sun during part of the day. 

The puparia were usually close beneath the surface of the earth, and the evidence 
suggested that the maggot had wandered but little from where it was dropped. 
Under each log was usually harder ground, interspersed with softer pockets where 
babblers (Crateropus) or small mammals had previously scratched. The puparia 
tended to be congregated in these pockets, but they appeared in some cases to give 
evidence also of gregarious settling by their mothers. 

I carefully identified all logs, old or fresh, under which puparia were found and 
was most interested to note that the very great majority of them were of two species, 
chiwhanga and mukarati (Ormosia angolensis and Burkea africana), which were by 
no means the commonest trees in the woodland. It was not that the fly chooses 
these out, for all evidence (and there was a great deal) showed that it goes after 
feeding to the nearest shelter, in some cases very diminutive. The reason was that 
these are actually the commonest logs on the ground, and they are so because they 
are the most durable timbers present and also do not readily burn. Some of these 
logs had quite likely lain for twenty or thirty years, very gradually dwindling, whereas 
a fallen Brachystegia may disappear in three. The eventual exploitation of these 
areas will result in the early clearing out of these two useful trees (the Ormosia for 
fencing posts) and the consequent confinement of the fly for its log breeding places 
to species that are attacked by insects and burn. This is one of the various facts 
that show how concentrated human activity assails the fly from several different 
directions. 

The localisation of the breeding centres in this area suggests that the possibility 
of attacking the fly on the lines suggested by Lamborn might require consideration. 
Some of the logs at the Kanyezi vleis that were cleared by me of pupae at the end of 
my first visit (early July) had already many fresh pupae under them when I came 
back at the end of the month, where hartebeests had been present. Breeding was 
going on both in June and July and in some quite cold weather, but emergences 
from the pupae secured only began taking place in numbers with the advent of 
really warm weather in early September, and continued into November. The 
pupae, like those of Lepidoptera, will apparently stand a small amount of damage. 
Out of 15 that I put aside as damaged in the finding on 30th July two afterwards 
produced flies. 


Glossina brevipalpis. 


I obtaimed no live pupae of brevipalpis, but found a few batches of pupa-cases 
in the Brachystegia bush on the Buzi east of Spungabera, one of which contained 
nearly a hundred puparia. In the case of this fly the batches were beside lyimg 
places of bufialos and wild pigs, and the flies at the moment of dropping the 
maggot had in the biggest find (Pl. xiv, fig. 1) evidently been resting for the 
most part on the undersides of the coils of large rough lianas (Cissus, Landolphia 
and Bauhima), apparently favourite resting places. Some of the puparia were 
found under logs end some of the replete gravid females taken during the bush-pig 
incident (p. 337) were resting on the lower surface of fallen Uapaca sansibarica 
(737) D2 


366 C. F. M. SWYNNERTON. 


stems, under which the pigs had been lying. The ground (shale) was in two cases 
exceedingly hard and compact, but humus was present in the situations in which 
the pupae were found. There seemed to be no such localisation of the breeding 
centres as I found with morsitans. 


Glossina pallidipes. 

I have referred to the puparia of this species elsewhere (p. 346). They were 
taken with those of G. morsitans and under the same logs, but in very small 
numbers indeed. None were found west of the Sitatongas. 


Glossina austeni. 

I have already given some details of this species (p. 348). The very distinctive 
puparia were all under four logs. One, more or less rotten—with only two 
puparia—was beside a game path used by bushbucks and waterbucks. Another 
with 30, showed many blue duiker “forms ” about it , but these were too recent ; 
leaf-carpet does not preserve traces well. The third, with 23 puparia, was also in 
dense shade and exactly at the junction of the primary-type forest and some heavy 
Markham bush (mubfeya). Right up to the log, in the latter bush, were the old 
tramplings and lying places of elephants. A similar log in similar shade lay 18 
yards away ; 1t was away from the tramplings, etc., and covered no puparia. The 
fourth log is described sufficiently elsewhere (p. 349). 


XV.—Net EXPERIMENTS. 
Resting Surfaces. 

When studying morsitans, and, later brevipalpis, I erected (Pl. xv, fig. 2) a large mos- 
quito net, 9 feet high, over cut-back tree-trunks, shrubs, etc., and furnished it 
with stems having different kinds of bark, stones, etc., in order to study the resting 
habits of the flies. These I turned into the net in some numbers—well over a. 
hundred in the case of brevipalyis. Some pallidipes were also included in each 
experiment, and each was continued for from three to four days. None of the three 
species used confined themselves to the few feet next the ground; they settled, 
colour conditions being correct, up to the full height available. I have noticed 
unconfined morsitans resting up to at least six feet from the ground. 

The rough-barked stems were selected in preference to the smooth, and large 
and small holes in the trunk and grooves in the bark were freely utilised for hiding 
in. A completely smooth-barked trunk was entirely neglected. Diplorhynchus 
mossambicensis was a favourite with the large tsetse, and a distinct colour-harmoni- 
sation took place also, the blacker tsetses (morsitans) choosing blacker bark, the 
greyer grey bark, the brown oncs (brevipalpis) brown bark and the underside of 
rough or knobby lianas, on which they easily passed as one of the knobs. 

For the morsitans experiment, in pyrophytic wooding in which the trees always 
show, on one side especially, blackening by the grass fires, the Diplorhynchus trunks 
were specially selected on account of the strong contrast between the colours of 
their two sides. Shade conditions were about equal (as the result of overhead shade) 
and, the position of the branch being also suitable, the female flies in particular 
always tended on settling to select the blackened side. On this side, furthermore, 
they settled chiefly where the black of the raised cork ridges alternated with pale 


THE TSETSE PROBLEM IN NORTH MOSSURBISE. 367 


brown grooves and this was at 3-4 feet from the ground. Below this area the bark 
was uniformly black, above it rather more uniformly pale end uncharred. The 
females also settled, the colours being right, mainly on the lower side of a branch 
or log, the males settling as frequently on the upper sides. A raised stone and 
clods of earth were also used for resting under in the brevipalpis experimznt. Leaves 
and thin twigs were used by active, not resting flies ; these flies were readily dis- 
turbed. The majority of the brevipalpis remained quiet during the heat of the 
day, but began to move freely and to buzz against the gauze at sunset. It was 
then chiefly that this fly, including its females, settled appreciably on non-protective 
surfaces—leaves and twigs, etc. Those definitely at rest on protective surfaces 
on the other hand, or in holes, were hard to flush, some (females, and particularly 
more or less gravid or replete females) even allowing me to tap on the bark beside 
them. The excellence of their concealment was shown when I finally cleared the 
net. Both I and two smart natives searched every inch of its contents most tho- 
roughly, as we thought, until we could not find another fly. I then passed my 
hand over the various surfaces and in this way flushed four more flies, all femé les, 
from the Diplorhynchus trunk. It would seem that Bagshawe’s suggestion—that fly 
does not haunt shady papyrus on account of the contrast in coloration—may well 
be correct. 


A certain number of flies were generally to be found on the net and restless ; 
these were always males. When the net was cleared and the flies examined one 
by one this observation was fully confirmed. The hiding flies were mostly, but 
by no means entirely, females. From my observations in the field also, it seemed 
elear that in Glossina, as in so many other animals, the female trusts mainly to 
eoncealment for defence against enemies, the male more largely to activity, and 
that the difference in the requirements of the sexes and their methods of meeting 
them is the chief reason for-the female’s special seclusion, though Lamborn’s factor 
(avoidance of males) may be operative also. The smallness of the meals usually 
taken by the males in the field—commonly mere snacks—bore a relation no doubt 
to their need for activity ; not so much to relation to enemies, for the male flies 
were quite capable of hiding, if necessary, but in relation to trevelling and the cap- 
ture of females. It seems certain also that the males need for food must be far 
smaller than that of the female. The results generally suggested the consideration 
of a contributory method of fighting the fly which I shall refer to below (p. 379). 


Distance of Attack. 


Another point tested was the distance from which brevipalpis and pallidipes 
will attack passing animals. It was undertaken at the end of my stay and I was 
unlucky in the matter of wind, light variable airs prevailmg during the main 
experiment. However, rushes to the side of the net next the cattle occurred at 
6 and 8 yards distance, and once, just before sunrise, at 18 yards, the shrub growth 
that blocked the view on this occasion showing clearly that it was a matter of 
scent. When the net was to windward of the cattle no general excitement took 
place, though a very few flies might fly towardsthem. Theshort distance from which 
attacks by brevipalpis took place in the sunny hours was illustrated in my clearing 


368 C. F. M. SWYNNERTON. 


experiments also. If I was right in believing that in the experiment on the Inya- 
marimu (described fully on p. 373) the individuals of pallidipes that attacked 
the cattle did so from outside the clearing, the small number that came to the cattle 
compared with the considerable number of these flies outside suggested that about 
30 yards may be the extreme limit of this fly’s attack under favourable conditions. 


A live goat tied inside the net attracted little attention. The flies bit occasionally, 
but allowed themselves to be driven off easily, and none fed to anything like 
repletion, yet the two oxen, when brought close up to windward, always attracted 
a great rush, even on the part of females that had been resting quietly on protective 
bark surfaces and had not been in the least attracted by the goat. I should say 
that I was myself much more readily attacked in the net than the goat. The 
liver of a cow was also not specially attractive. 


XVI.—NATURAL ENEMIES. 


During my first week at the Kanyezi vlei I was camped 150 yards or rather 
more from a considerable cluster of male flies. These were preyed on daily,* 
sometimes continuously and for a long time, by half a dozen to eight birds belonging 
to three species—Dicrurus afer (African drongo), Bradyornis ater (mimetic fly-catcher) 
and Bradyornis murinus (mouse-coloured fly-catcher). From the continuous 
nature of the attacks, and the immense number of house-flies (Musca domestica) 
that tame drongos of my own have shown themselves capable of devouring in a 
very short time, there could be very little doubt that the birds were making con- 
siderable inroads into the male fly population, and it was likely that the practical 
disappearance sometimes of this cluster was in part due to them. The birds perched 
on the low trees bordering the vlei and dropped to the tsetses in the grass below. 


In my general experiments I found that many birds disliked Muscid flies, including 
Stomoxys, replete or empty (actual Glossina not tested), but that drongos, fly- 
catchers, stonechats, swallows, bee-eaters and some commoner small searching 
birds (Crateropus, Phyllastrephus, Apalis) liked them much ; drongos and swallows 
(certainly also fly-catchers) continued to feed on them when so replete,as to refuse 
all non-Dipterous insects. The digestion in at least two of these last three groups 
is so rapid that (in my experiments) even when the birds were replete three or four 
minutes’ rest made room for two or three or more flies. These birds must be rela- 
tively formidable enemies to a fly the males of which display themselves so freely, 
and it is a pity that the population of such enemies must be limited by the dry 
season food supply. I have sometimes seen swallows hawking continually after 
insects on grass-stems and grass-blades, picking them off while flying, so that 
male morsitans is a likely enough object for attack by these birds. Birds are in 
the habit of paying special attention to insects, otherwise acceptable, that are present 
in numbers together, so that in localities in which male morsitans is being kept 
scattered by the presence of much game it must lose a far smaller proportion of 
its population through the attacks of birds. 


— = 


* The presence of the birds at this spot was noted practically daily ; they were only 
continuously watched on about two days. 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 369 


Craleropus kirkt (Kirk’s babbler) and the two guinea-fowls, Gullera edouardi 
(crested) and Numida mitrata (Kast African), were observed once or twice (out of 
many times) scratching in positions like those in which tsetses deposit their larvae, 
as was also the bristle-necked bulbul (Phyllastrephus capensis).* A more interesting 
observation was made on an elephant shrew (presumably Petrodromys tetradactylus) 
ia dry but shady donga entering the Buzi at the south end of the Sitatongas. We 
had searched for and found puparia (austen) under a large fallen tree and, going 
on to search under other logs and stones, found we had been anticipated by a small 
animal. For a distance of 47 paces along the donga every hiding place that would 
have been used by tsetse had had the surface soil scratched out and the spoor and the 
droppings were, I should say (and the natives also were certain), distinctly those of 
one of these elephant shrews. 1 have actually watched the above elephant shrew 
in the wild state scratching thus for slightly buried insects and, having kept indi- 
viduals in captivity, am tamiliar with their droppings, etc. 

The natives stated that ox-peckers (Buphaga) regularly prey on tsetses that are 
feeding on game animals, and that from the movements of the birds it is obvious 
that the flies employ against them the sidelong movements I have referred to in 
relation to attack by knife and assegai. The flies also no doubt continually come 
within the ken of the birds that carefully search tree-trunks and twigs, and it is 
no doubt in relation to these that their choice of protective resting surfaces is of 
use. Such birds form a very large proportion indeed of our small bird population. 

Spiders of various bark-haunting species that hide in crannies and leap on their 
prey accounted for a number of the flies I had under observation in the net, and it 
is probable that these are the female tsetse’s most.important enemy. One tsetse 
(morsitans) was seen caught in the net of a web-making spider. Dragonflies on 
several occasicns in fly country inspected the backs and necks of my companions, 
A wasp once did the same. On one occasion, travelling from the Buzi to the 
Mtshanedzi, five brevipalpis were under the cattle, four being under the leading ox. 
We ran into some dragonflies, which at once began passing to and fro immediately 
under the belly of that ox. I saw no captures, but these four brevipalpis had dis- 
appeared when the dragonflies left us. The one survivor travelled with us for over 
one and a half hours and remained about our camp for long afterwards. Dragon- 
flies were exceptionally and extraordinarily numerous in 1918, but the tsetse popula- 
tion remained large. I do not remember seeing any ASILIDAE. They only become 
abundant here in October. 


XVII.—TsEtTsE CONCENTRATIONS. 


My closer observations on this subject, which may be of high practical importance, 
were limited to a single morsitans centre, but I searched and studied much country 
in which there was no concentration, both before and after the commencement of 
leaf-fall, and secured much native information also. I obtained, | believe, a fairly 
clear idea of the conditions obtaming in this particular fly-area. 1 spent a fortnight 
in all at the Kanyezi fly-centre. This was a large and very definite breeding 


* The existence of soft pockets of earth, breaking the harder crust under most of the 
logs examined, constitutes indirect evidence also of the scratching of some animal or 
animals in these places. 


370 C. F. M. SWYNNERTON. 


concentration in country relatively devoid of the fly. Secondly, my native expedition 
reported coming across several such concentrations north of the Lusitu, with the 
same male crowds at the edges of similar vleis, the same ease in finding puparia and 
the same relative dearth of tsetses in the similat country between. They brought 
me both morsitans and pallidvpes and puparia of the former. Thirdly, the Masando 
vleis and some vleis east of the Mapapa (see Map) were both stated by the local 
natives to be the sites of concentrations, how great I do not know, and to have been 
so in the present season. I had myself found a concentration at the Masando in 
1900, but in 1918 we found the flies on the grass of the vlei-edges only in parties 
of three or four—as at Kanyezi’s, when a male crowd had been dispersed. Buffalos 
and elands were present in small parties. We were led to the more easterly vleis 
by a native (subsequently in my service and apparently reliable), who stated that 
he had seen the flies there in great numbers only a week or two before. I had no 
cattle with me and only two flies were taken, but a large herd of buffalos had arrived 
meantime. These two sets of vleis and Kanyezi’s are all within a very few miles of 
each other. 


Leaf-fall Concentrations. 

When I left Kanyezi’s on 1st Augsut it seemed likely on the whole, though 
uncertain, that this scattered, unconcentrated fly was even somewhat sparser than 
it had been in June. The few that the cattle picked up on that particular day were 
at vleis or glades—not that this was unusual. Natives on the Umvuazi River had 
reported an increase in the flies about their villages, and one way in which the insects 
might travel thither appeared to be illustrated by the fact that the flies that were 
attaching themselves to us left us (one of them after following for 54 miles), not in the 
rather barer bush, but on our reaching the somewhat shadier wooding beside streams. 
The natives stated that this tendency to collect at the streams becomes more marked 
rather later, but not (Kanyezi said—and I think we saw) at the expense of the vlei 
population, which also increases. 

The natives attributed the streamward movement of the fly in the later dry 
season to the fact that the general drying up of the country forces the game thither. 
“The int-hesi then congregates there to feed on the game.” While this explanation 
is not quite accurate, it is certainly to be judged from what I have said above that game, 
or natives, either going to the streams or merely travelling across them, will tend 
to deposit there at this time any flies that they may have picked up either in the 
drying-up country generally or at vleis, etc. These Jeaf-fall (and soil-moisture ?) 
concentrations, whatever they may amount to here, are evidently to be dis- 
tinguished from the more permanent breeding centres, although these also will 
doubtless receive accessions from outside at this time. 


Primary Concentrations. 

That the concentration of the fly in the last-named centres is not purely the result 
of dry season concentrations there of the game seemed certain. There was no 
concensration whatever of the game at Kanyezi’s vleis, where I stayed a fortnight in 
all and at which fly was plentiful and breeding, nor had there been since the rains. 
‘There was even less of a concentration at the Masando in June 1900, when I found 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 371 


tsetses there in great numbers and very troublesome, but could find nothing to shoot. 
The concentration (in this case merely the arrival of a large herd of buffalos) at the 
more easterly Mapapa vleis in 1918 took place when the flies were (according to the 
natives) already abundant there, and the small parties of buffalos that were appar- 
ently keeping down the numbers of the visible fly at the Masando in 1918 could 
hardly be described as a concentration. It was certain from the detailed distribution 
of the large number of muddy puparia taken at Kanyezi’s vleis that the fly con- 
centration was already in being here in the late rains, when also a great buffalo herd 
was temporarily present ; and as my flies and pupae were indubitably largely the 
descendants of the depositors of these puparia, it might be said that fly concentration 
found in the dry season may be the result of game concentration either then or 
previously... Even this (as accounting for the presence of a fly centre, apart from 
the mere question of its population) was negatived by the native statement that this 
fly concentration is permanent though variable in numbers, that it had been there 
before the buflalos came (as it had also survived their departure), and that the 
animals had stayed a very few days only. In testimony to the great size of this 
herd my guide, Kanyezi, who lives near and had seen them, pointed out strong paths 
that had been beaten out and vleis that had been heavily pitted with spoor in a single 
day. The natives said that the other centres seen or heard of by me were also present 
each dry season, though again with a varying population. The conclusions that I 
would draw from all I saw and heard, including the tact that away from these centres 
we failed to find any but very stray pupae, even when bufialos and other game had 
been abundantly present, are that what Shircore has called “ primary centres ” 
undoubtedly exist; that their actual (not their apparent) population no doubt 
varies with the immediately preceding food supply; and that in their situation 
beside vleis they have hit off the best obtainable combination of the three desiderata 
of food, moisture and shade (for vleis, generally speaking, and their neighbourhood 
are attractive to game and other animals, and the wooding bordering them is 
quickly back in leaf). It is even possible (in an area in which morsitans happens to 
be dependent on the larger mammals) that a kind of selection is continually taking 
place by which those vleis that for any reason have become neglected by the game 
for a considerable period cease to be fly centres. It was certainly strange that 
Gundoda’s vleis, well watered, midway between Kanyezi’s and the Masando and 
quite near each, and in the same wooding, uncleared, but carrying a human popula- 
tion and a more permanent shortage of game therefore than the others, should have 
been shown by a fairly exhaustive examination to be practically devoid of pupae 
and to have produced few flies. Some of these few, latterly, were marked flies 
released by me in the Kanyezi vleis, and it is likely that the great number of flies 
said to be at Gundoda’s in the rains come first from this and other vlei series. 


Village Concentrations. 


A concentration of another kind than the two referred to above, and presumably 
one of a very shifting nature, is that which appears to take place at native villages, 
even in the rains, and particularly whenever the fly is abundant. Gundoda (whose 
village, besides affording good shade and being situated at the edge of a well-watered 
viei, is a tempting dismounting-place for fly) {described to me well how in the 


372 C. F. M. SWYNNERTON. 


spring and early rains, as they would be sitting in the shade by the water, 
attended it might be at first by a merely reasonable number of tsetse, every 
newcomer or passer-by would bring with him an accession of flies till their numbers 
became unbearable. Some would go again with departures, more would stay, and 
they tended to increase in numbers from day to day. At that time every one, 
sitting or travelling, carries a leafy switch and uses it incessantly. The probability 
that the great majority of the flies of these village concentrations are males was 
suggested further by my failure to find puparia at Gundoda’s. 


Leaf-fall in relation to Concentration. 


It would seem that in one of the three forms of concentration I have referred to, 
leaf-fall must be a highly important factor. That there are other factors producing 
concentration is however certain; for on my first arrival in the morsitans country 
in June, concentration at the vlei centres was already in full operation, yet leaf-fall 
(the season being late) was not yet noticeable. The same would apply to my visit 
at the end of May and beginning of June in 1900, when also high concentration 
was present. When I left on Ist August 1918, leaf-fall was in many places pro- 
ducing a visible effect, and it is possible that this may have brought about the 
apparent increase in the numbers of pallidipes at the Kanyezi morsitans centre. 
Otherwise things had not, to all appearance, been so very different in June from 
what they were in August, Whatever the factors that bring about these “ primary ” 
concentrations, the possibility is suggested that measures against them might 
not have to be confined to the very brief period of maximum leaf-fall. All would. 
really depend on the value of the “scattered ”’ fly for the local survival of the species, 
and I am inclined for the moment to suspect that this may be greater in the case 
of pallidipes—everywhere seemingly a rather scattered fly—than in that of 
morsitans. 


The Kanyezi centre was certainly, to judge from its puparia, a focus of morsitans 
rather than pallidipes, and it is conceivable that these primary centres may be a vital 
point in this fly’s social organisation. 


The rural population of brevipalpis, as I saw it, was much heavier and more 
evenly distributed than that of morsitans and therefore less easily distinguished 
from its rather more populated patches, in the primary thickets. In a less generally 
suitable area (such as the granite-gneiss actually is) or in a season (if such occurs) 
which really stripped the Brachystegia wooding of the “ Oblong,” it would doubtless 
undergo leaf-fall concentration. 


XVIII.—TxEe Recent OUTBREAKS OF NAGANA IN MOSSURISE. 


The fly, driven from a large piece of deciduously wooded country by Umazila’s 
measures, never greatly abandoned its permanent haunts except on the Sabi and 
in the cleared portion of Gogoyo’s, the Mwangezi, etc., and within 7 or 8 years 
after the relaxation of the measures in 1889 it was found re-occupying its old (and 
its present) wet season quarters in so far as bush conditions and travelling facilities 
then allowed. Since then, within my own observation, the wooding of the heavily 
deciduous areas hes increased greatly, and areas close to the British border that 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 373 


18 years ago were relatively bare are now very fully wooded and capable of giving 
a temporary support to tsetse. 

Up to a very few years ago cattle were still kept successfully in Mossurise within 
a few miles of the British border from the Puizisi to Maruma, and from Spungabera 
to the Inyamgamba. These, except the Spungabera cattle, which have suffered, 
have been largely wiped out by successive attacks of nagana, especially during 
the last three years. At the same time isolated outbreaks, here and there, have 
taken place on farms just on the British side of the border. In a remarkable pro- 
portion of cases a connection has appeared to exist between the outbreak and the 
wanderings of buffalo. 

The incursions of buffalo into the cattle-carrying areas have undoubtedly increased 
greatly in the years concerned. The northern outbreaks coincided with or followed 
the invasion of Mafusi’s country by considerable numbers of buffalos from the 
south and their coming in contact with the brevipalpis-pallidipes area there. 
In the section opposite Spungabera (across the Buzi) these animals are certainly 
far more in evidence, and in far larger numbers, than they were some years ago, 
as I know from shooting trips on the Upper Mtshanedzi in 1901 and 1902. 

Elands have also greatly increased, as have pigs; and it is likely enough that 
the fly itself has increased within its permanent haunts in consequence of the great 
and progressive increase in its food, and of the earlier burning of the grass. This 
would contribute to the menace, and it is by no means incompatible with the fact, 
of which I am convinced, that such destruction of the big game as might be easily 
feasible would not alter the fly’s permanent boundaries, though it would certainly 
prevent it from being carried into contact with the cattle. 

The great increase in the wandering game has undoubtedly been the main factor 
in the outbreaks of the past few years. But it may be said that it is especially 
hard to distinguish the separate original infections where natives’ cattle are con- 
cerned, owing to the likelihood of infection by mechanical transmission by other 
biting flies that is mcurred through their placing the survivors in the herds of 
their neighbours, and by their movements of cattle in connection with their 
matrimonial affairs. 


XITX.—CLEARING EXPERIMENTS. 


These were carried out only to ascertain the effect on the fly of the clearing of 
undergrowth (PI. xvi). 

Experiment |. Jo ascertain the effect of clearing the undergrowth in primary forest. 
The whole width of the dense undergrowth of the Inyamarimu forest strip was cleared 
to a length of 80 yards. The width varied from 40 to 70 yards. It was incidentally 
open to penetration and passage from the sides by any flies there might be in the 
Brachystegva-Uapaca bush on either side of it. At 11.30 on the following day the 
two oxen were driven into the middle of the clearmg. They carried in with them 
one fly, a male brevipalpis picked up in passing near some of the uncleared bush, 
and this was at once removed. They were stood in the centre of the clearing and 
from time to time moved across it a little though not to its edges. They had been 
placed many times before in this spot and had never failed to be attacked by large 
numbers of brevipalpis. This time in three hours only four came to them. 


374 C. F. M. SWYNNERTON. 


G. pallidipes and Labanus had never before been noted on them in this dense 
wooding, though they occurred in the bush outside. Yet in these three hours nine 
pallidipes (all but one females) and 31 Tabanus were taken on them. 

I then turned the two oxen just mto the uncleared undergrowth alongside and 
they were at once attacked by brevipalpis. My collector captured 15 on one of them 
here in fifteen minutes and no pallidipes or Tabanus were seen at all. 


The effect of clearing the undergrowth alone in banishing the big fly was, I think, 
well demonstrated. It is perhaps more likely that the other flies were attracted 
to the cattle from the bush outside than that they had taken up thei abode in the 
forest as the result of the clearmg of the undergrowth. Tkis would suggest that 
attacks of pallidipes in the sunny hours are undertaken from a greater distance 
than those of brevipalpis, bemg equivalent rather to the latter fly’s evening attacks. 

Experiment 2. To ascertain the width of undergrowth clearing necessary to protect 
a strip of road from attacks by brevipalpis in sunny weather. In Brachystegia bush on 
the Buzi, the cattle in passing along a certain track four times in all had on each 
occasion been attacked by brevipalpis all the way along from the thickets passed 
closely or passed through. I cleared the dangerous undergrowth, but none of the 
overwood, for a paced length of one mile and a width of 25 to 40 yards with the 
track in the middle. TI left three small points not fully cleared—some small Diplo- 
rhynchus trees with low-hanging branches and Bauhinia, a small primary-type 
thicket, and a piece of uncleared sapling growth. I put the cattle through and they 
were attacked at the first two of these places—nowhere else at all. I brought them 
back just outside the clearing and they got flies all the way. In the afternoon, 
having cleared these spots, I put them through again, bringing them back this time 
along the track. They got no flies at all either way until driven out beyond the 
cleared strip at each end. They were then each time attacked at once. 


It was to be expected that this narrow clearing would be of use only in sunny 
weather and this point was tested later on an overcast day. The cattle were driven 
along the cleared mile and 32 brevipalpis were taken at them, coming in all the way 
along. They still failed to be attacked here in sunny weather. 


Obviously then, very little clearmg is needed to safeguard a given piece of road 
from this fly, provided that the cattle pass along it in sunshine. A considerably 
wider clearmg would be needed to render it safe at all hours and in all weathers. 


Clearing by Arsenite of Soda. I had previously carried out successful experiments 
in poisoning with arsenite epiphytic figs in forest on my own property. This sug- 
gested its use in clearing for tsetse, for the difficulty in ordimary axe-clearing is 
that the stumps at once (and repeatedly) send up fresh growths and form coppice 
that is even more suitable for the fly than the growth cleared. A recent examination 
of some of my experiments shows that the strength of an ounce of arsenite to a 
pint of water (the maximum strength tried) is insufficient to be generally effective, 
though it killed some trees and inflicted damage on many others that may enable 
the fires to burn them out. I hope shortly to be in a position to state the best 
strength for this sort of work. Given this and the amount of liquid required per 
acre for particular kinds of bush, one can estimate very rough'y the cost of this 
kind of clearing. 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 375 


Mere ring-barking—with an upward-pointing frill—is required instead of cutting 
down and this will economise part of the extra cost of the arsenite. * 

The balance of extra expense will doubtless in particular cases be justified by the 
fact that the work will not have to be repeated, but it will be seen that where great 
areas of dense undergrowth are concerned this will be considerable. Naturally 
cattle should not be run where arsenic has been used in the last few months. 


X X.—MEASURES FOR THE CONTROL OF T'SETSES IN MOSSURISE. 


The parts of the Mossurise district which are more especially suited to white 
settlement are the two dolerite areas (a) that between the Lusitu and the Mtshanedzi, 
and (b) the Gwenzi country from Spungabera to Mount Sngunu. These areas would 
split up into a very great number of 1,000 hectare farms, practically all good. The 
soil is for the most part rich and particularly adapted to Arabian coffee (a good close 
settlement product, if the labour should be sufficient), the natives raise fine crops 
“even when crops are more or less of a failure elsewhere, the rainfall is considerable. 
and reliable, and a good deal of local labour is available ; the grazing throughout 
is good and the grass early. It was pitiable to see my bait-cattle being followed for 
miles like a circus by the children of some of the villages we passed in a country that 
should be carrying its thousands of head. There are good permanent streams 
everywhere. 

If there should be an unwillingness to sacrifice the Mafusi rubber forests—which 
will always carry fly and be a menace—there might still be a large number of farms 
in the remaiming area. If the outlet to a market is not through British territory, 
there is, for produce, the Lusitu-Buzi waterway investigated by Snr. Roma Machado 
and capable of being opened as part of the settlement scheme. The eventual 
creation of a safe winter route for cattle along the south of the Buzi is also not 
improbable. f 

The Umzila results, already described, show clearly—and it cannot be too much 
emphasised—that settlement properly planned will protect itself. As settlement is 
bound to be the eventual policy, no matter how long deferred, it cannot be said that 
the ultimate future of the more delectable of the Mozambique Company’s infested 
areas is necessarily seriously compromised by the presence in them now of fly. 
Umzila’s results even suggest that some day in the very far distant future the question 
will be settled by the natural increase of the now protected native population. 
“Properly planned ” settlement in fly will not consist in the giving out of isolated 
farms, scattered over the face of the country. The failure to keep cattle at the 
Dysart Concession is a case in point. 

There must be a definitely planned settlement scheme, affecting a large block 
of country together, on some sound agricultural basis. The closer the settlement 
can feasibly be the better, and first and foremost amongst the conditions of occupa- 
tion must stand the effective clearing of the less freely deciduous types of woodland. 


*T have tried boring, both by auger and by down-slanting strokes with a narrow 
native axe, but find that this leaves unaffected areas of bark between the holes and kills 
only particular branches, the poison (as might be expected) taking effect in a vertical, 
not horizontal direction. 
ft Later information suggests that the Sabi even now offers a safe route. 


370 Cc. F. M. SWYNNERTON. 


These will be inferred sufficiently from what I have said in this report, but I will add 
here that in the piece of country in question, along the streams and rivers and in 
larger blocks on the Lusitu, first-class timber trees—mahogany (Khaya), musando, 
mowana (Adina) and others—are present in quite unusual numbers. These, floated | 
down the Lusitu, might in some cases help much to defray the cost of clearing. 

Secondly, no ingress of large game must be allowed from areas still under fly. 
The best barrier—with the elephants away—might be a strong, patrolled fence ; 
but if the fringes of the area are sufficiently closely settled it is likely, even from our 
present imperfect success on the lightly settled border, that this fact alone, with 
shooting, will suffice to keep off the elephants, buffalos and elands. It is these three 
animals that probably chiefly matter. 

Umzila’s principle—the settling and clearing of the low-lying guard-area only, the 
enclosed hill mass then taking care of itself and being perhaps disposed of later at an 
enhanced value—would be well worth consideration and investigation. Under such 
a scheme, carried out with thoroughness, it seems at present fairly certain that cattle 
could, after a few years, be kept safely and in numbers on the dolerite. 

If, on the other hand, the settlement should have to be a gradual growth from 
small beginnings, its safest base would be the deciduous part of the British border, 
a block at a time being settled and special measures being taken against the buffalos. 


The settlement of the two permanent fly-areas themselves would mean the end of the 
menace. The great strip on the Lusitu that is now occupied by rubber forest, both 
east and west of the river’s southward turn, represents some of the finest agricultural 
ground in the country and 1s in immediate touch with the Lusitu-Buzi waterway. 
Much clearing is necessary, but the farms here could be small. The “ Oblong” 
offers greater difficulty. The soils on the whole are distinctly useful, though less 
rich than in the other brevipalpis-area, but there is relatively little permanently 
running water away from the big rivers. Pools, however, persist in most cases and 
water should be obtained in this formation at no great depths by boring and sinking. 


X XI.—Discussion oF GENERAL METHODS OF TSETSE CONTROL. 


Game Destruction. 

In view of the evidence I have already alluded to (p. 336 and elsewhere) we are 
bound to keep an open mind with regard to the possibility that even G. morsitans 
might survive the destruction of all large mammalian life. Yet the fact that under 
present conditions game-paths are the regular rendezvous of the sexes, and that 
the connection with game appears generally to be an essential point in their rather 
complicated social scheme, makes one hesitate to reject the alternative view. 
Lamborn (Bull. Ent. Res. vii, 1916, p. 37) has argued well for the view that it need not 
be a large game population that will support fly, and this might be a sufficient 
explanation of Umzila’s continued trouble in the rubber forests. 

In a relatively clean-stemmed area like that on the granite-gneiss, in which also 
bush-pigs are not over-abundant, it is to me very conceivable that wholesale game 
destruction might banish the fly. An obstacle, however, would be that the whole 
territory is one vast game area, so that the game would pour again into a given 
section of it as soon as the persecution was relaxed—unless an effective barrier 


THE TSETSE PROBLEM IN NORTH MOSSURISE. <a is 


were created such as could probably only be made a permanency with settlement 
behind it. Whether the returning game would bring fly in again would depend on 
(1) whether the whole continuous fly area had been cleared of fly, or (2) whether 
the portion cleared had been split off from the uncleared portion by an effective 
barrier against the fly itself. 

West of the Sitatongas the country is much more jungly, and wholesale game 
destruction is proportionately more difficult. In addition this country abounds 
in bush-pigs, which are difficult to destroy and which in anything approaching 
their present numbers can probably alone support the fly, with baboons, abundant 
cane-rats and other animals which may all contribute to its sustenance. I consider 
that it will be impossible to starve the fly at all generally by ordinary game destruc- 
tion here, at any rate before the country is very fully settled, though buffalos and 
elephants might be banished by adequate and persistent shooting. 

What can be done is to protect particular places, like Spungabera and the British 
border, that are outside the fly and are threatened only by the wanderings thence 
of the bigger game. Fencing, the judicious placing of native kraals and shooting 
are amongst the possible measures, and organised and repeated drives might be 
undertaken locally. I have suggested similar means of keeping the fly from being 
carried into any areas that may in the future be settled. 

Again (cf. p. 355) it seems to me that game destruction should check the advance 
of a fly-belt, whatever its effects on the ground already infested. No problems 
are at present better worthy of study than these—(1) the extent to which tsetses 
will travel independently of game, and (2) the distances that the female will travel 
on game. 


An Attack on the Concentrations. 


At first sight, the weak point of morsitans in the area in which I have seen it would 
appear to lie in its most striking attachment to the vicinity of more or less moist 
spots and in the great numbers in which it breeds there. One is strongly tempted 
thereby to suggest that for areas in which this is the rule further experimentation 
should be carried out on these distributing, and probably receiving, centres ; 
experiments in local clearing of varying widths, in planting vleis and their margins 
with the heavy wooding avoided by morsitans,* poisoning and gassing the male 
clusters (torpid on dewy mornings) and the bush just immediately adjacent, catching 
with the aid of cattle, pupa-destroymg or any other methods against either sex 
or either stage however remotely promising, suggest themselves as worthy of trial 
before the idea of attacking morsitans at its concentrations is finally given up. 

The essential point to ascertain, however, supposing such measures to be capable 
of exterminating the fly at the concentrations, is the value or otherwise of the 
“stray ’ flies (which comprise both sexes) for the survival of the species, and whether 
these would continue indefinitely to escape the measures applied at the centres— 
always supposing that, in the area concerned, they are not and cannot be driven in 


* To check radiation, introduce shade conditions inimical to the fly and spoil the game’s 
early grazing, also if the trees that succeeded were of a definitely ‘‘ drying’’ type, like 
Eucalypts, to help to dry the ground. Such a measure, even if locally successful, would 
doubtless be too expensive to use except under special circumstances. 


378 C. F. M. SWYNNERTON. 


completely by leaf-fal. or simultaneous burning. It would be difficult to ascertain 
their value until one had already successfully applied the local measures. 

There are other difficulties also to be faced in dealing with the individual centres. 
Their probable large number in a given area—though in the area visited they 
occupied only a small number of the places that appeared suited to them—gives rise 
to difficulties (1) in relation to finding them all, (2) in relation to the number of 
white workers that it might be necessary to employ when simultaneity seemed 
important and time short, as in work that might be planned against the emerged 
flies. Here the period when emergence is already rapid, but the fly not yet greatly 
scattered, might be of special importance. There is also the fact that the game 
must first be banished from each centre before the flies will present themselves for 
treatment in anything approaching their full numbers. 

Broader lines of work are undoubtedly preferable if any should be found effective, 
and it will be most interesting to see the result of the Southern Rhodesian Govern- 
ment’s present measures against morsitans, planned, as I understand they are, 
on game removal and large measures of bush destruction. 

To discuss one or two of the points, nevertheless, it may be said that the great 
reinforcement of the visible fly sometimes called out by the presence of my cattle 
showed that ordinary collecting by hand is a very inadequate method, especially 
with game about. Collecting round cattle would be far better, but their length 
of life under continual attack is a point to be considered in relation to the financial 
feasibility of the scheme. It might well be prolonged by antimony injections, 
but would otherwise (to judge from my experience) be a matter of a very few weeks 
indeed. Better still would it be if the flies coming to the cattle could be automati- 
cally poisoned. The effect of frequent arsenical spraying, although it seems quite 
unlikely to be useful, might be tested experimentally, for it has not been my experience 
that a tsetse keeps its proboscis buried to the bulb all the time it is feeding, and 
a possibility I have referred to elsewhere (p. 341) might also be worth a trial. 

If the breeding concentration seen by me at the Kanyezi centre should be typical, 
it suggests that in areas where such concentrations are the rule it might be possible 
to devise useful measures against puparia. Artificial breeding places, unless 
scattered nearly as freely as.the present fallen logs, though with a regularity that 
would make them more easily found, would not, I think, be useful ; for my indirect 
evidence all pointed to the probability that the heavily pregnant female does not 
fly many yards from where she last fed. 

The present situation at Gundoda’s vleis suggests that the placing of Kafir villages 
at the fly-centres might break these up through the effect on the game, but with 
so many apparently eligible sites vacant the concentrations would presumably 
reform elsewhere. This measure alone would only be useful if it were possible to 
plant a population at such places through the area generally. 

So far as G. brevipalpis is concerned, the observations spoken of on p. 351 would 
require to be greatly added to before we could assume that this fly’s permanent 
boundary could be put back merely by the splitting up of the bush on its western 
margin. The point is worth looking into further. In any case the game would 
require to be destroyed or excluded in order that the fly might not be carried back 
in the rains. 


THE TSETSE PROBLEM IN NORTH MOSSURISE. ‘379 


Assistance to Natural Enemies. 
Natives destroy great numbers of birds, in both the adult and tisadiion Pras 


but I fear it would be impossible to make protection effective. It perhaps matters 
less here, as the native population is in parts not great and in the “ Oblong ”’ it 
is almost non-existent. The same difficulty applies to elephant-shrews which 
are greatly trapped. It is a pity that these useful animals cannot be protected, 
the more so as they are put to death in a peculiarly cruel manner. 

Rewards for the skulls of small carnivora brought in might be useful in better 
populated fly areas, probably not here. There would in any case be a doubt whether 
the natives, who trap freely for these animals already as articles of food, could be 
stimulated very much more by a prospective reward. The greatest enemies. of 
nestling birds here are the tree-snakes (Dispholidus) and probably the only effective 
remedy against these lies in severer fires. 

Indirect assistance to birds would take the form of destroying trees, etc., the bark 
of which is specially protective to the tsetses. This method is inapplicable to 
morsitans and pallidipes because so large a proportion of the trees in Brachystegia 
bush have suitable bark, grey and somewhat charred, that wholesale clearmg would 
be necessary. In a brevipalpis area the destruction of Diplorhynchus mossambicensis 
and all rough barked lianas would probably be distinctly useful and should form an 
essential part of any hand-clearing scheme ; not but what other species I could 
specify are of probable use to this fly in varying but usually less degree. 


Protection of Stock and Riding Animals passing through Fly. 

I have already described my clearing in this connection (p. 374). In addition I 
found that very frequently the. animals were not attacked if the carriers 
were walking immediately ahead of them, without an interval. I sometimes 
walked at the head of the carriers myself and then noted flies (brevipalpis) coming 
out to me and turning back—rejection, evidently, after inspection. | 

Again, when I divided the cattle into two lots (two in front, one after an interval 
behind—I never had more than three) I found that the first lot drew off all the fly, the 
last animal commonly getting none. . 

In the following experiment both these principles were called in. I wished to 
protect a donkey in passing through the heavily infested three mile strip following 
the Inyamarimu. I placed two oxen in front ; the four goats followed them at from 
forty to eighty yards behind ; then (just behind) came myself and a native, then the 
donkey with its rider, a native walking in each side of it and one behind ; next, at 
some distance, came a cow. The oxen both carried fly all the way and were some- 
times assailed by many ; the goats’ driver captured a brevipalpis on himself, but the 
goats escaped entirely ; one only came to the cow. On emerging from the dangerous 
bush, the oxen were turned sharply into the grass and took their flies with them, the 
other animals, which now passed ahead, getting none. The control experiment, 
if such were needed, had been provided two days before when, on the same path, the 
donkey was sent on a detour to avoid a bad bit of forest. It ran into a thicket and 
sustained quite an attack and had to be brought back behind the cattle. 

This suggests that a mob of cattle might be protected in travelling by placing the 
less valuable animals a little ahead. Naturally a mob of cattle is a bigger mark 

(737) E 


380 —. Cc. F. M. SWYNNERTON. 


than my animal or animals that marched second, but that relative and perhaps 
nearly complete protection would be afforded I am convinced. 

I have heard it stated at second hand from a man who had had some experience 
of running cattle through fly that 1t was chiefly the outside animals of a mob that 
were attacked. It may be possible to prevent even this. 

I failed to carry out any experiments in the matter of spraying or wiping animals 
with paraffin or other deterrent liquids, though it had been my intention to do so. 
Against brevipalpis the animals’ lower surface should be sprayed. | 

When biting flies are abundant it 1s a common experience to have them attack 
the cattle at once on their emerging from an arsenical dip. An internal tsetsefuge 
might, however, be experimented for and be used where the number of cattle was not 
too large for treatment. During our first epizootic of African Coast fever, in 1900, 
when I was already losing stock seriously, I tried dosing the surviving cattle each 
morning with a double handful each of garlic, crushed and given in a bottle of water— 
as we had then no dipping tanks. I am inclined to attribute the fact that I saved 
a much larger percentage of our small herd than did any of our neighbours (who lost 
nearly everything) in part to the deterrent effect of the garlic on the ticks. I do 
not know its effect in relation to biting flies, but it might be tested, and if it were 
successful, garlic-eating might become a useful if obnoxious habit in sleeping- 
sickness areas. 


Clearing Measures. 

(1). Clearing by cutting down. Where not followed up, the good effect of this 
measure must be very transient, for the stumps quickly produce coppice of a type 
that is specially liked by pallidipes (PI. , fig. 46)—or, under shade, by brevipalms. 
Where, on the other hand, clearing by cutting down is followed by very late annual 
burning, its effect, very generally speaking, is likely, when the grass is sufficient, to 
last as long as the late burning is kept up. It is said that a large proportion of 
Uapaca cut down during the rainy season does not grow again. I have no observa- 
tion on this myself beyond the fact that Uapaca trees are more easily killed in native 
gardens by burning than are various other trees. The reaction of each of our 
commoner tree species to the cheaper methods of clearing would be worth ascertaiming 
by special experiments. 

(2). Clearing by arsenite. Experiments that I carried out on pieces of wooding 
selected as representing an average of a type that was well infested with brevipalpis 
did not suggest that the clearing by arsenite of such continuous Brachystegia bush 
as fills the “ Oblong” and runs north to the British border past Spungabera would 
necessarily run to a prohibitive figure for this class of work, though the full strength 
to be used remains uncertain. Naturally the clearing of great areas is In any case 
certain to be expensive. 

Superintendence and transport of water, ete., have to be considered. While 
native police might superintend the actual working groups, generel supervision by a 
Kuropean would be necessary both for the efficient organization of the water trans- 
port and of the work generally, and because it would be unsafe to leave large quanti- 
ties of poison in the sole care of native police. The best effects from poisoning would 
probably be obtained when the sap is up, that is in the spring and summer. Water 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 381 


would doubtless give much difficulty in some localities in the spring months, but 
the country generally has water in streams or pools. 

It will be gathered from this that poisoning offers certain difficulties in addition to 
its possible high cost. It could be put into effect most easily and cheaply round a 
settled station like Spungabera, where there are also mere patches of wooding to be 
dealt with ; but here the greatest care would have to be taken to keep all stock from 
treated areas until all possibility of their obtaining poisoned leaves had passed. 
The method would be still more useful to occupiers of farms under a settlement 
scheme in the fly, for cattle would not yet be present. 


Killing by arsenite, followed by a course of late fires, should be exceedingly 
effective in removing woodland, and the finality of it, with the fact that it requires 
vastly less labour than the other radical method, stumping, would be worth the 
expenditure that a considerable increase in the strength of the poison might entail. 

A local point to be remembered is that my observations were carried out in a year 
in which bush generally tended to carry leaf in the dry season better than usual 
owing to the heavy preceding rainy season. Brachysteyia wooding has this tendency 
at all times, except when exposed to relatively rigorous conditions, but if it should 
be found later that in certain dry seasons the Brachystegva-dominated bush of the 
“ Oblong” generally does become too leafless to harbour brevipalpis, the clearing 
of the ravine-type thickets might suffice—if at any time in the future the hand- 
clearing of this area comes to be considered. 


(3). Clearing by judicious native settlement. The first lesson to be drawn from 
Umzila’s results se2ms to be that we can make a seasonal fly-area safe by broadly 
(perhaps narrowly) clearing its margin and preventing passage of game. 

In the permanent fly-area and portions of another permanent fly-area that he 
cleared of fly the result was brought about, apparently, mainly by bush destruction, 
though game also was greatly reduced. 

It is perhaps a pity that the humane methods of the white man make difficult the 
wholesale transportation of populations for the elimination of tsetse areas, even in 
the interests of the population in question. Our methods would be less wasteful 
than Umzila’s. We should begin with a close botanical and oecological study, and 
we could produce Umzila’s result with a half, a quarter or an eighth of the force used 
by him. 

On a small scale, with such an inducement to settle as the remission of their tax 
(meaning much to the native, and to the Government a great saving in the cost of 
clearing by other methods), the plan is still very feasible. When examining the 
Spungabera problem I was struck by the impression already made in a single season 
by the native Umgazaza on a piece of wooding that constitutes one of the special 
dangers of the place. With a few more Umgazazas settled here this piece of wooding 
would go—at a small cost to the Company. The soil to be cleared is rich, the natives 
would raise good crops, and they could dispose of their surplus grain a mile away 
at Spungabera. Other wooding on the Spungabera hill could be treated in the same 
way. 

The pity is that the cultural operations of our natives do not include the taking 
out of tree-roots. If, however, they left all stumps standing—and they leave a 

(737) E2 


382 Cc. F. M. SWYNNERTON. 


very great many in any case—these, in secondary bush, could be treated with arsenite 
by the owners themselves under the supervision of police, who might be sent round 
with it for that purpose. This should render the clearmg permanent—at a cost. 
Surviving primary-forest stumps are killed by the grass fires after the cessation of 
cultivation. 


(4). Clearing by regulated burning. Of the probable good effect of this measure 
I can speak with an assurance that is based on nearly twenty years of interested 
observation and experiment; interested, because my own land has been in question, 
and I have had to study there the effect of early, late and no burning on pasture and 
afforestation respectively. 


I have already (p. 325) contrasted the effect,of early and late burning on thicket 
and sapling growth. The following quotation is from a paper of mine, read in 1917 
and published a year later, and written with no thought of tsetses in my mind. 


“The fact is,” I wrote (S.A. Journal Sc., June 1918, p. 16), “that land from which 
fire is excluded tends to go back to dense bush. Even the more open grass-veld here 
is full of stumps that seldom get further than a one season’s shoot. The very fires. 
that have rendered their existence on that ground possible, by driving the [primary] 
forest off it, keep them from growing up until, some year, a poor burn, it may be, or 
no burn, allows of a second season’s growth being superadded to the first and gives 
a more fire-resisting bark to the latter. ... Keep the fire from such a piece of 
ground—or burn too soon—for several years, and these shoots grow up and eventually 
in places become so dense as to reduce the grass and the severity of the fires and to 
allow semi-forest types as Markhamia lanata and Albizzia chirindensis to spring 
up amongst them, as I shall describe below, and eventually to replace them. The 
result [even] when this occurs so far from high forest as not to obtain seeds from it, 
is a form of dense thicket.”” On my own land Uapaca-Brachysteqia areas burned 
regularly and more or less late have failed to develop appreciable undergrowth, 
while areas which, for winter grazing, have been burned irregularly and early, have 
developed it (PI. xvii, fig. 2). The latter areas, which were clean-stemmed when I came 
here,would now carry G. brevipalpis ; the former would not. Elsewhere in the irregu- 
larly burnt area new coppice wooding has sprung up in open ground from long 
suppressed underground stumps and roots and could already shelter pallidipes. 


The converse statement—that late fires will destroy already existent growth—is. 
true also. It is particularly true of a late fire following a year in which burning: 
was omitted ; and I have sometimes been much struck—the last instance was late. 
in 1917—by the large size of the pyrophytic saplings that such a fire has killed down. 
On occasions even large pyrophytic trees are definitely killed by it, at any rate in 
such a season as 1913. Of the effect on primary-type thickets I have given an 
instance on p. 325 and have seen further instances myself. 


With regard to the statement that late burning was a practice under the Zulu 
regime, I can say from personal observation that there was less wooding, and less 
undergrowth under existing wooding, in the Northern Mossurise district even when 
I lived and travelled in it from eighteen to nineteen years ago than there is now after 
a further period of unregulated burning. It is likely that this probable result of 
their late burnings contributed much to the Zulus’ success, particularly of late, in 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 383 


the margins of the “ Oblong,” and it is unfortunate that such fires should necessarily 
have ceased at the very moment when whole districts of native gardens were 
abandoned to grow up into coppice growth. | 

The effect on reproduction of burning at a time when a large proportion of the 
trees are in flower must also not be overlooked. I wrote on 5th December 1906 
on examining a late “burn” between Chibabava and Arucate and observing 
that the flower crop generally had been destroyed, ‘it will be interesting to note to 
what extent this late burning will effect the seeding, and consequent replacement 
of dead individuals.” There can be little doubt that it will affect it considerably 
in the lower types of wooding. 

Late burning is no emergency measure against fly. In some areas especially, 
where the grass is already well reduced, its full effects may take long toshow. Again, 
its abandonment will mean the return of the old conditions, for the underground 
stumps and roots of pyrophytes are extraordinarily tenacious of life. It must be 
regarded as a piece of administrative policy and be kept up annually. That its 
adoption is necessary I am quite convinced, if only for the reason that the country 
generally is at present reverting more and more to wooding and becoming increasingly 
fitted for tsetse, even in those areas in which the cover has hitherto been sparse and 
light. The recommendation of the measure, in face of its having to be kept 
up indefinitely, must be that the fires take place in any case and that the annual 
cost of regulating them will be trivial. The latter may even be met by the fines 
imposed on unauthorised burners, and the actual burning, being done at a signal 
by the kraal natives themselves, will cost nothing. 

Apart from the prophylactic value of the measure just mentioned, its effect, grass 
conditions being equal (which they are not), should show first and chiefly in relation 
to G. brevipalprs and (in the area in which this fly depends mainly on coppice) 
pallidipes. It will not affect the grown pyrophytic trees in any wholesale manner 
{unless in a very exceptional season or unless they are weakened by ring-barking). 
Its effect in relation to morsitans might even be regarded as problematical, for that 
fly and pallidipes in the same wooding appear to be independent of undergrowth. 
The systematic burning back of smallish growth will assuredly tell as the established 
trees pass maturity and eventually go, with nothing to replace them, but the time 
involved will be long. 

The objections that may occur should be discussed. One is that the loss of the 
wooding at springs may dry these up. Many of the springs rise already in open 
vleis, and wooding on damp ground (including big Khaya and Adina) is in any case 
not easy to destroy by burning, except occasionally in such a season as 1913. But 
better an occasional spring lost than a continuance of the tsetse; and the Zulu 
clearings, sufficient to remove the fly, do not seem to have caused any shortage of 
water. At worst, occasional lost spring-heads could be allowed to revert to wooding 
after the tsetse had been eliminated. The exposure of the ground to wash by burn- 
ing shortly before the rains is not an effective objection, as it has been our experience 
here that the growth of spring-burned grass rapidly overtakes that of winter-burned 
grass, which itself affords little protection to the soil in October. Loss of, or damage 
to, the secondary wooding generally—unless of the very finest Albizzia-Milletia- 
Pterocarpus monsoon types—need not be regretted, for reasons which I have stated 


384 . C. F. M. SWYNNERTON. 


in the paper from which I have already quoted. It comprises types that are 
relatively useless from every point of view. As for rainfall, the merits of wooding in 
bringing about this kind of condensation are in dispute, and at best only the big- 
wooding area of the Lusitu would need consideration ; but all local evidence goes 
to show that it is the mountains, not the wooding, that perform this service for us 
here. The objection in any case is applicable to any clearing measures. 

Another difficulty is administrative. Even on the British side of the border, 
where unauthorised fires are forbidden by legislation, such fires still take place. 
This will happen also here, but the local set-backs involved wil] weigh lightly in the 
balance against the benefit that should accrue from the general success. The penalty for 
unauthorised burning should be severe, and even if it should be necessary to employ 
a few extra police-boys for four months in the year, the measure would be a very 
cheap one. There should be no discontent over it amongst the natives, as late 
burning represents their own old custom, and, whatever their infringements, they 
still speak of it as the correct method. Dr. Lawrence, speaking of the fact that 
native huts and granaries must have the grass round them burned at an early date 
to protect them against the grass fires, has suggested that it will be more difficult 
to protect against late fires, and that it will also be difficult, when one of the guard 
fires gets away, to discriminate between real accidents and pretended ones. It 
seems to me that the real accidents and cases of hardship will be reduced to a mini- 
mum—along with the carelessness or lack of adequate hoeing preparation that 
produce most of them—by a refusal to excusethem. ‘The natives adapted themselves 
successfully to the late burning of the Zulu regime and even now have sometimes 
to meet late, fierce fires, as they had to in some of Gwenzi’s heavy grass-jungle 
country in 1918. Naturally they would receive ample warning. 

Another difficulty that has been raised in conversation—but it is no difficulty 
at all—is that late burning is supposedly bad for the pasture. The excellent grasses 
left us here by the Zulus, who burned late and regularly, are a sufficient reply. It is 
true that continued burning checks the development of a “close sole” of still 
better grasses that otherwise follows on heavy stocking by cattle ; but if you stop 
proper burning before you stock, you Jose your pasture altogether. The land goes 
to dense wooding, as it is now fast doing everywhere in Mossurise. Wholesale 
late burning 1s In any case not being recommended for country that is already 
carrying cattle. 

Simultaneous burning off of the infested country—and more or less simultaneity 
would in any case result from postponement—would probably discommode the flies 
on the wing in a more direct fashion. It would banish the game for the time being, 
and in the case of brevipalpis in the “ Oblong,” it would probably force the fly either 
to take refuge in the ravine-type thickets which a course of late burnings should 
eventually destroy, or, in part at least, to perish. Its probable effect on morsitans 
seems more doubtful in view of this fly’s relative independence of shade, but 
observations by Lamborn (Bull. Ent. Res. vii, p. 39) and others show that it is usual 
even for this fly to desert burnt areas. 

Unemerged pupae would no doubt remain in any event to repopulate the area, 
but Lloyd took a greater percentage of dead pupae from burnt areas than from 
unburnt ; and if I am right in my expectation that the heat of the October fires 


THE TSETSE PROBLEM IN NORTH MOSSURISE. 385 


will reach a considerably higher proportion of pupae than those of the earlher fires, 
the anticipated effect on the flies on the wing would be well worth obtaining also. 
Birds, again, as I have shown already, probably attack tsetses most when these are 
in numbers together, and the temporary clearing of the game from a large area will 
remove the agency that keeps the smaller tsetses scattered, while hunger and the 
black background should increase the flies activity and visibility. 

The one thing that has made me hesitate to recommend simultaneous burning 
is a slight doubt as to its exact effect on the insectivorous bird population itself. 
October burning in any case encroaches somewhat on their breeding season, but not 
to an important extent—that is, the early nesters that have had their nests destroyed 
have still plenty of time before them, and even in ordinary years an amazingly large 
proportion of them has to make a fresh start as the result of the depredations of 
enemies that are likely to be adversely affected by late burning. Again, a fire and 
the few days following it are feast days, for a very large proportion of the winged 
insect population escapes it and, being then more conspicuous against the universal 
blackening, is more readily detected by the birds, which may sometimes be seen. 
searching the burnt ground in parties. The real point is whether the wholesale 
destruction everywhere of the insects’ own food will so much reduce their numbers 
as to react adversely on the birds. I think that there will be no difficulty here at 
all, for in this district at that time of the year the response of the vegetation after 
burning is immediate and most rapid, and a burn gets repopulated with wonderful 
speed, the new brood of grasshoppers even showing colour adaption to the scorched 
surroundings ; but the effect of simultaneous burning on the insectivorous birds 
should be noted nevertheless. _ 

In short, the effects of stmultaneous burning, while very promising, have yet to be 
accurately observed ; but the good effects of late burning have actually been demon- 
strated. 

I wish, therefore, to make it the one outstanding recommendation of this report 
that late burning—with, very frequently indeed, a year of no burning—should be 
given a trial over a considerable number of years ; also that, in the first season at any 
rate, an effort should be made to have the burning as simultaneous as possible over 
a large block. The rubber forests and finer Lusitu wooding generally could, if 
desired, be left out of the scheme. Early burning thereabouts, every year, will 
save them. It will also unfortunately spread both them and the fly. The fine 
timber there must be considered also; and against the reduction of the cost of 
clearing which late fires might have effected by the time this section comes to be 
settled, must to some extent be placed their destruction of the surface humus. 
Elsewhere this is being destroyed in any case by the useless fires ; it may as well 
therefore be destroyed by useful ones. The very deciduous areas at the higher 
elevations need not necessarily be included either, but they are not sufficiently 
extensive to be worth excluding, especially as their wooding (for the most part of little 
value) will remain a great annual danger so long as any permanent fly survives below. 

To give the measure a definite experimental value sample plots or areas in each 
type of wooding should be selected, described, charted and photographed. These, 
examined from time to time, would afford a measure of the success or failure of the 
experiment. 


ri 
he Te a ha 


wel ys ripe! iF ee 


ie 


Sad ds es when ‘ome : 
tr; » 


¥ 


Pe ll 


- : 
rs Es 


(ee 


er 
‘— pidly 


ay 


LE 


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etal ‘> 


Buti. ENT. RESEARCH. VoL. XI. PART 4. PLATE IX. 


Primary ‘“‘rain” forest; the large trees are African mahogany (Khaya nyasica’. 
At suitable elevations this type of forest harbours Glossina brevipalpis in the 
dry season. 


BuLc. ENT. RESEARCH. VoL. XI. PART 4. PLATE X. 


Fig. 1. Highly deciduous secondary forest on Mount Umtareni in 
August, with Pterocarpus angolensis and Terminalia sericea. 
Capable of sheltering Glossina brevipalpis and G. pallidipes in 
the wet but not in the dry season. 


Fig. 2. Lowland bush-savannah on the basalt at the younger 
Gunye’s, Gombretum dominating; harbours the overflow of 
G. morsitans from the Brachystegia for much of the year. 


BULL. ENT. .RESEARCH. VoL. XI. PART 4. PLATE XI. 


7 


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3 


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$ , P ~ 
BMG 


A typical scene in the Madanda rubber forest. An Erythroxylon-Landolphia 
invasion of tree-savannah and savannah-forest, capable of harbouring 


G. brevipalpis and G. pallidipes. 


a 


met Sipe aed Met = 


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PLATE XII. 


PART 4. 


XI. 


VOL. 


ENT. RESEARCH. 


BULL. 


In Uapaca-Brachystegia bush; two shade- 
layers, and G. brevipalpis present. 


Fig. 2. 


In Uapaca-Brachystegia bush; lower shade- 
layer only, therefore no G. brevipalpis present. 


Fig. 1. 


Buti. ENT. RESEARCH. VOL. XI. PART 4. PLATE XIII. 


Fig 1 In Uapaca-Brachystegia bush; only an upper shade-layer, 
and therefore no G. brevipalpis. 


Fig. 2. Landolphia wooding of the Lusitu rubber forest type; with 
a lower shade-layer of this high, very dense type G. brevipalpis 
was present even when the higher trees were leafless. 


+ 


T) 


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Butt. ENT. RESEARCH. VoL. XI. PART 4. PLATE XIV. 


=a 


aN 


t . J 


ss 
2 


Fig. 1. Nearly 100 puparia of G. brevipalpis were taken under this 
fallen Piptadenia buchanani at the spots indicated by the papers. 


Fig. 2. A very typical breeding-place of G. morsitans under a 
long-fallen trunk of Burkea africana. 


— oo —"” a= 


-, — 


BuLt. ENT. RESEARCH. VoL. XI. PART 4. PLATE XV. 


Fig. 1. 


Site of a male cluster of G. morsitans; the short grass was 


very green and luscious, and the flies occurred both on it and 
on the longer grass near, 


Fig. 2. 


Experimental net and its furniture, as used for G. brevipalpis 
and G. paillidipes. 


~~ 
a a 


aes als 
| J A 


PLATE XVI. 


RESEARCH. VoL. XI. PART 4. 


ENT. 


BULL. 


uncleared ; 


Undergrowth 


G, brevipalpis present in abundance. 


Clearing experiment No. 1. 


Ile 


Fig. 


; G. brevipalpis 


has entirely disappeared. 


The same spot with the undergrowth cleared 


Fig. 2. 


Butt. ENT. RESEARCH. Vot. XI. PART 4. PLaTE XVII. 


Fig. 1. Coppice type of secondary bush, strongly favoured by 
G. pallidipes west of the Sitatonga Hills. This bush has grown 
up in 12 years on deserted native gardens. 


Fig. 2. A thicket on a piece of land belonging to the writer 
resulting from early and irregular burning for a good many 
years; it was previously clean-stemmed wooding. 


But. Fees: R 


SO —— 


=~ ae 


Explanati 


ostly sparse and low, bush-savannah, ‘*Umpoza” dominating; fly 
Breen areas.- 


ther poor but + continuous Brachystegia savannah forest with ma 
ans and, in places, a few brevipalpis. 


posure— sandstone, shale quartzite and good Brachystegia—Uapa 
rs. 


for the most part highly deciduous; permanent fly absent. 


wooding chiefly of the various dense-secondary types, especially fer 
a brevipalpis) more localised then than in the rose and dark-gre 


nd Umtshanedzi. - 

ts. Wooding contains an unusually large element of the primary ( 
Pteleopsis. Much Landolphia kirkii in North. -Brevipalpis appare 
Tleleopsis Pangolpnia Kirkil Brevipalpis 


~—~~~~~ —Route of two native expeditions. 


pproximately, but distances (cyclometer and timing) probably nearl 
nce. 


ps. s.f. = secondary forest. f.p. = primary forest-patch 


o Portuguese frontier and numbered beacons. _—_ 
New kraal below Chipungambira forest. F. = Ferry and kraals of 


: From ‘1" north-eastwards brevipalpis was present almost conti 
ojd *‘5"' flies (pallidipes) seen on June 13th, 1918. ‘6° a combi 
&/uzi. 8" brevipalpis, do, +. “9” Jacks high-elevation pallidip 
e, 1900. “12°' and ‘*13"’ most westerly and southerly morsitans, 
idipes than Jack's were taken on the Inyamadzi. 


le-keeping under Umzila and Gungunyana; also, roughly, the line 


2-keeping now pushed back to the old boundary of the pre-Umzila 


Alleged 
qe es. @ e__ Recent nagana outbreaks. 


Fly. matic and gives merely a fairly correct general idea of the prese 
map and the portions (taken by compass from frontier beacons) o 
ply fairly correct also the windings of the Buzi from Itamba northw 
and the positions of some Kafir kraals and river-crossings. 


O06), showed low country to S.E. covered densely with trees, stated 
parse. : 


Burt. Ent. Researce. Vou. X1., Part +- 


Chimanimani 
Massif 
(Peaks over 8000 feet) * é i 


Y ab 
aE ———=—- SAMBA 2" here, Je 


J 
os reported pa Pticulary 


ven roe torent N eae 
Ps és} oe (a, MUCHAMBA y Wee” 
Mou 5 rath cs wh ¥ — no se \5 m. To "Buzi Col “ 
nt Singwekwe : F: INYAMSIZWA_7,ru R- a Go BH ence’ saihing boat te Heine 
or Pene, 6000 ft. 3 Mee zs pe ; lg et CHIBUYE ‘ \ _7. \Chirenda 
: Z 
Y, : e 
GUDZA NAY io Seatarel SNE se? fram eh S 
: 7 ‘rom Chironda for a doz 
1 Mag patcres SPS > Honecat lrren progeaten Seopa uareR arta 
. fiers! busts) 3 Scattered thickets (largely Pavetta galigna). Further 
: es vast more open, but with bigger trees (still open) 
7 reported here Be near river. Approaching Imyamita (196 ny. from 


Guzt Colonial) more open still, 
BECC Pe scattered palms ang 


Map to Illustrate 
REPORT ON A STUDY 
of the 


J : ye 
SS Marowery. H 


Hh = Makwakwa Y ES 
YEW: e s> 
NORTH MOSSURISE A Moyet (and elder Gunye wae 
3Gurdoda : §Mangunde 
7) Mapapa vieis, etc. 1 


TSETSE PROBLEM. 


<a Fly Ppenenms nid very 

& SS eer Fe troublesome in! 
SSH asarans Mee eL 
wl 


VitoFre.\ MEPLORE ty HS 
pe ea 25 
rp! a 4 ~~ om 
- 6, b 3 e hy 

‘ “yee Fea S _-—> 2 Chibabava 
- Subse “%, A Ce ‘\ 
Baayaberive rarer fe 
“i Ms To Arucate and 

u R@asanande net Madanda forests. 


" 
MO and snene 
Let iew! ane ger! 
BY pas: “WIPE ance, “HOS ee 
\ 


pare Bont 
mwwrncaNe,, BE caSe 
Sec 


ie case 
U 


more prominent 


points of a somewhat broken 


° 
5 f - 
es] Mainly basalt; wooding mostly sparse and low, bush-savannah, “*Umpoza” dominating; fly pi pes and morsitans, apparently chiefly as overflow from the salmon and 


7 
7 
Pole fkrasl” . 3 1 PS wo (pallidipes) dark-green areas. 
(ae ficest piece of country ; MSE a Pp: s pi é : 
heh ; , a F eu OS™ Ot. Dos =] ainly granite-gneiss; rather poor but + continuous Brachystegia savannah forest with many vieis. 
Spunfebiea® be 5 oe Soe cae ranhe ~ FAO » Fly pallidipes and morsitans and, in places, a few brevipalpis. u 
‘ YN owana * eo Fi 5 
Dengedza f.p- (6) BLONG Rr > We BY eo pene A great sedimentary exposure—sandstone, shale quartzite and good Srachystegia—Uapaca savannah forest, continuous brevipalpis everywhere in the thickets, pallidipes 
4 / with ari aecos small hillS) d Playyena ach srallerinura Bere, cs 
f ee | Mainly dolerite; wooding for the most part highly deciduous; permanent fly absent. 
<4 
ae BA Mainly dolerite; broken wooding chiefly of the various dense-secondary types, especially fern-entwined. In the main deciduous, but late in the season, and the permanent 
a27 ZA fly (pallidipes ana brevipalpis) more localised then than in the rose and dark-green areas, Brachystegia (and fly) on scattered Sedimentary exposures, Ramturartesy 
©, between) Puizise and) Umtshanedzi. a a 


Mainly dolerite and schists. Wooding contains an unusually large element of the primary (chiefly ravine”) and the finer secondary “monsoon” types « Khaya, Piptadenia 
TET Bon AU IEEh any Acacia caftras tn Albizia, Milletia, Pteleopsis. Much Landolphia kirkii in North. -Brevipalpis apparently abundant throughout, pallidipes present. eee 
i & reticulata, Albizzia hypoleuca Kigeli 


Bik EN not dangerous all the year an een Route of main expedition. ~ -————— — Route of two native expeditions. 
wee’ © . . « 
ees 28 POG: a Buzi route: shown very approximately, but distances (cyclometer and timing) probably nearly correct. The notes appended to it were made in 1906-7 and the wooding may 
ey iter RO ogee have increased since. 


Four main working camps. s.f. = secondary forest, fp. = primary forest-patch 1. = G. austeni vleis, 


7 ao . o—~o—o 1 
ADEE rah oo e806 gh” Anglo Portuguese frontier and numbered beacons. “——_ _ Buffalo passage areas, routes and approaches. U, = Umgazaza’s kraal, 
ZZ; oe Soo Corse oy 2 Ma. = Mapondo's. N. — New kraal below Chipungambira forest. F. = Ferry and kraals of Siyaghiya anal Shilling. wis u 
7 (Cn s 

Ts ee bi ASebosher™ord © O.- Flies taken furthest west. From ‘1" north-eastwards brevipalpis was present almost continuously in numbers, "2" — a fly taken by Mr, Jack. ‘3" by Dr. Lawrence, 
pushy.an If UmbIya Ne. ee ae August. “4" and “5” flies (pallidipes) seen on June 13th, 1918. “'6" a combined attack by pallidipes and brevipalpis, 7 = brevipalpis continuous in thickets 

. i} ater from here to Buzi * brevipalpis, do. +. “9” Jacks high-elevation pallidipes, August, 1917 " = brevipalpis dry season limit here. ‘11 Wilder an 

25 } 2 Lawrence's tsetse. pT “13 "" most westerly and southerly morsitans, respectively. * = highest elevation brevipalpis, 3000 ft. or somewhat over, 
“10 i Fan-palms — Hyphane — More westerly pallidipes than Jack's were taken on the Inyamadzi. ‘ 
yj) become common from id ae 


| here eastward, throughout. Eastern boundary of cattle keeping under Umzila and Gungunyana ; also, roughly, the line of maximum clearing in the Puizise-Mtshanedzi area. 


oGungunyana’s, 2 
capita Kay great numbers — LE 
ver ~~ Much Bauhinia 99S> ~ 
© AS HLIT = = =====- Recent boundary of cattle-keeping now pushed back to the old boundary of the pre-Umzila period 


vane cS 
ZINYUMBO er 


Hot Diplorhynchus (Miatawaypeonumon— in paris Note___. The contouring is diagrammatic and gives merely a fairly correct general idea of the presence of hill-masses and their relative importance. The frontier is taken from the 

Brinn gmecmian i fran sin unos Ss faontoiChimbiys official Rhodesian map and the portions (taken by compass from frontier beacons) of the Sitatongas, higher points of the oblong, Umtarene and the Spungabera 
details are probably fairly correct also the windings of the Buzi from ltamba northwards and, in a more general way, the course of the Umtshanedzi and the entries 
of the tributaries and the positions of some Kafir kraals and river-crossings- 


Alleged 


. Some definite Zulu failures. @ e-_ Recent nagana outbreaks. 


_ View from here (Nov, 1906), showed low country to S.E. covered densely with trees, stated by natives to constitute  Maguasha," or dense forest, that along Buzi route was 
seen to be very Sparse. 


XK 


Bale & Danielsson 4 


387 


MOSQUITOS COLLECTED IN PALESTINE AND ADJACENT 
TERRITORIES. 


By Capt. P. J. Barraup, F.Z.S., F.E.S. 


These notes include observations made on the mosquito fauna of Palestine, for the 
period July 1919 to August 1920. 

In September and October 1919 a somewhat rapid tour was made through Syria 
and Cilicia, while from time to time short visits have been made to Egypt. 

The climate of this part of the Mediterranean region is divided into wet and dry 
seasons, the latter usually prevailing from May to October, inclusive. In the northern 
parts of Syria and Cilicia the dry season is of rather shorter duration. From 
November to April there is a heavy rainfall over the larger part of Palestine and the 
countries to the north, but in the lower Jordan Valley, and in the deserts of the 
south, the amount is much less. The winter is mild, especially along the coastal 
belt and in the lower Jordan region, the thermometer seldom falling to freezing point. 
In some seasons there may, however, be a considerable snowfall in the mountains. 

With the advent of summer, the country quickly becomes arid. Rivers and 
streams are few and insignificant, and the inhabitants of the towns and villages 
are obliged to rely for the most part upon wells or supplies of rain-water collected in 
catchment tanks. In certain parts of the country there are, however, perennial 
streams, and in some cases these form extensive marshy tracts, affording breeding 
grounds for mosquitos during the greater part of the year. Irrigation is practised 
throughout the dry season in districts where the supply of water is sufficient from 
springs or surface wells. 

The summer weather in the Jordan Valley can only be described as tropical. 
Along the coastal plain there is usually a sea breeze, but when this fails, the moist 
heat is often trying. The higher parts of the mountain ranges enjoy a succession of 
hot sunny days, tempered by cool breezes, with a greater fall in the temperature at 
night. The maritime plain, consisting for the most part of rich grain lands, orange 
groves and olive yards, varies in width from a narrow strip where the mountains 
approach the sea, to many miles where they recede. 

The mountains of Judea and the Galileean hills, rising here and there to 3,000 feet, 
form the backbone of Palestine proper. They consist chiefly of barren limestone. 
To the eastward they descend in tumbled masses to the Jordan Valley, a unique and 
stupendous crack in the earth’s crust, sinking to 1,200 feet below sea-level. On the 
far side the mountains of Moab divide the valley from the tablelands of Bashan and 
the Arabian desert. 

The larger part of Palestine, especially the central and northern regions, Syria, 
and Cilicia are malarious. A fairly high percentage of the native population in 
some of the towns and villages has been found to be infected, and well known 
malaria-carrying species of Anopheles abound. The duration of the malaria season 
roughly corresponds to that of the dry period, from May to November. 


388 CAPT. P. J. BARRAUD. 


In early summer the Bedouins, with their flocks, descend to the plains in search of 
pasture and water. From their habits it has been assumed by some authorities 
that the majority of these people are malaria-carriers, and in their wanderings assist 
in the spread of the disease, but I am not aware that this opinion is based upon any 
extensive investigations. It is possible that these people derive a certain protection 
from the fact that they are usually accompanied by numbers of the larger domestic 
animals, horses, donkeys, camels and cattle. I have observed that frequently some 
of these are stabled under one end of the curiously shaped bivouac tents used by the 
Bedouins. These conditions appeared to be favourable for the feeding and shelter- 
ing of Anophelines. 


Although I was unable to prove that Anopheline mosquitos prefer feeding upon 
animals rather than man in this part of the world, I have noticed, when searching 
camps, that they are usually to be found in larger numbers in tents pitched near 
horse or mule lines. In the towns and villages the number of large domestic animals 
would not be sufficient to supply food for the abundant Anopheles, and in these places, 
no doubt, they mostly subsist upon the blood of human beings. 


One of the most abundant domestic mosquitos of the country is Anopheles 
bifurcatus. I have seen it breeding in countless thousands in the basement rain- 
water cisterns, which are to be found almost universally beneath and around private 
houses. As this stored water often represents the sole supply for the inhabitants 
during the summer, anti-malarial work is difficult, and opposition is sometimes met 
with. 


In Jerusalem there are upwards of 4,000 basement cisterns, besides an extensive 
system of ancient drains and sewers, many of which have become blocked by 
subsidences. Some of these cisterns are of considerable size, having a rain-collecting 
platform as large as a tennis court, with a correspondingly large surface of water 
beneath. They are from 15 to 20 feet deep, and usually have only one opening about 
2 feet square, so that a satisfactory examination is hardly possible. Hven where a 
lid or cover has been made mosquito-proof, and a pump fitted to draw up the water, 
there is always an opening through which the rain enters. Mosquitos find their 
way in and out through this. Large surface wells protected by masonry are 
another frequent source of trouble. One of the most important medical problems 
with which the new Administration has to deal is the provision of an adequate piped 
water supply to all the towns, and the abolition of basement cisterns, surface wells 
and catchment tanks. 


In Egypt, where the incidence of malaria is much less, the three chief Anopheline 
carriers of Palestine, viz. A. maculipennis var., A. bifurcatus and A. swperpictus, 
are practically unknown. A. multicolor is there considered to be the principal 
carrier. It is common in the Canal zone, and especially abundant in the oases. It 
is in the last-named places that malaria is most prevalent. Here again the popula- 
tion is largely a nomadic one, tending to spread the disease from place to place. 
It is interesting to note that all the species of Anopheles so far found in Egypt occur 
also in Palestine, in spite of a wide strip of waterless desert separating the two 
countries. On the other hand, there are four or five species in Palestine which are 
absent from Egypt. 


MOSQUITOS GOLLECTED IN PALESTINE. 389 


Anopheles maculipennis, Myg., var. eu 


This is generally distributed over Palestine, Syria and Cilicia, but although 
recorded from Egypt by earlier workers, has not been found there in recent years. 


The adult has the wings less distinctly spotted, especially in the male, than the 
European form,* some specimens being unspotted. The resting position on a vertical 
surface is rather flat, this being more noticeable in the male. Gorged females 
resting on a tent roof hang down in the usual Anopheline position. 


Major S. R. Christophers discovered this faintly spotted form in Mesopotamia and 
wrote to me in regard to it, mentioning that the egg differs from that of the typical 
A. maculipennis (figured by Nuttall and Shipley in Journal of Hygiene, i, pl. 1, figs. 
1 and 2) in that it has a frill of air cells all round the edge, instead of one pair of lateral 
floats. This observation I have been able to confirm in the eggs of Palestine speci- 
mens. The full-grown larva and the adult do not, however, appear to differ structu- 
rally from the type, and it remains to be seen whether the variety here referred to 
should be regarded as a distinct species. 


The larvae are usually to be found in natural water both fresh and brackish. The 
adults have been found in very large numbers in tents and huts in camps, in various 
parts of the country, and often cause a heavy incidence of malaria. They will 
sometimes travel a considerable distance from the breeding-grounds, especially 
when emerging in large numbers. On 27th September 1919 I visited a camp at 
Toprak Kali, Cilicia, which was situated on a hill 500 feet high, and one and a half 
miles from the nearest water. In the earlier part of the year this site had been 
considered safe from a malaria point of view. The tents were, however, found to be 
heavily infested with both A. maculipennis var. and A. superpictus. Very similar 
observations were made on Ist October at a camp in the valley of the Ak river, near 
Marash, Northern Syria, from which large numbers of cases of malaria were being 
evacuated. In the first tent examined (square Indian pattern) there were over 300 
Anopheles in one corner, while a patient lying beneath had 70 inside his mosquito 
net. These were nearly all A. maculipenms var., with a few A. supermctus ; some 
of the former were noticed to be biting in the daytime. Although the river passed 
within a few hundred yards, very few larvae, or likely breeding-places, could be found. 
In marshy ground and rice fields at about one and a quarter miles distance, however, 
the larvae were in abundance. 


Breeding commences in April and May, according to the season, in the marshy 
areas along the coastal belt of Palestine. In the Jordan Valley, at depressions below 
sea-level, it probably begins earlier. During the first part of May 1920, I collected 
large numbers of larvae and pupae in a shallow brackish marsh lying amongst the 
sandhills, east of Haifa, and near the mouth of the Kishon river. Larvae of several 
other species, viz. A. multicolor, A. hyrcanus, A. mauritianus, Culex univittatus and 
C. tepuliformis, were found at the same time. By the end of May larvae were much 
less numerous, the day temperature of the water in the marsh having then risen to 
88-90°F. 


*The North European form has not been found in Palestine, but the two forms occur 
together in Macedonia. 


390 CAPT. P. J. BARRAUD. 


Adults of A. maculipennis var. were found in camps on the sands near the Kishon, 
and a few from time to time in camps above Haifa town at 300 feet ; also in the lower 
western part of the town, at least one and a half miles from the nearest breeding 
place, as far as I could discover. Larvae were never found within the town limits, 
either in covered or uncovered collections of water. 

At the end of April larvae began to appear in an area flooded by late rains, north 
of Acre town, in the neighbourhood of a battalion camp. This area soon became dry, 
but in the meantime large numbers of A. maculipennis var. had hatched out, and 
an outbreak of malaria occurred, lasting several months. The first primary case 
of benign tertian was reported on 28th May and of malignant tertian on 27th June. 

The species occurs from 1,200 feet below sea-level to 2,300 feet above. My collec- 
tion includes specimens from the following places :— . 


PALESTINE : Sarona (near Jaffa) ; Zummarin ; Athlt ; Haifa; Acre; El Afule ; 
Jenin ; Tabgha (north of Tiberias), along the course of the Jordan from the Sea of 
Galilee to near the Dead Sea. Syria: Damascus; Homs; River Ak Valley (near 
Marash). Crticta: Missis ; Jihan ; Toprak Kali. 


Anopheles bifurcatus, L. 

As mentioned previously, this is the most abundant domestic Anopheline mosquito 
of Palestine and Syria, breeding almost exclusively in basement rain-water cisterns 
and covered surface wells. I did not discover the larvae in any natural open water 
in Palestine, but at Beirut (Syria) larvae and pupae were taken from a small gently 
flowing stream containing much vegetation and over-grown with bushes. It appears 
to be essentially a cool-water species. The temperature of the water in basement 
cisterns in a coastal town, such as Haifa, varies between about 58°F. in winter and 
78°F. in summer. During the spring and early summer, when breeding is most 
active, the water was found to remain at about 62°F. Larvae may be found through- 
out the year in their favourite haunts, but feed up more slowly in winter than during 
the warmer months. The majority of larvae must pass their whole existence in 
semi-darkness, the sun’s rays seldom penetrating to these deep cisterns. 


Examinations were made at Haifa in N ovember, before the autumn rains, when 
many of the cisterns were dry or contained very little water. In most cases they 
were clean, with very little, if any, debris upon the floor. In others, which had 
probably not been cleaned out for some years, the walls had become green, and there 
was a thicker floor deposit. Further investigations in J anuary, after about six 
weeks of heavy rains, revealed the presence of very large numbers of adults. Collec- 
tions were made of individuals resting on the walls, within reach of the opening. 
At first I was under the impression that they were all females, but while carrying out 
a series of fumigation experiments, it was found that in cases where all the mosquitos 
had not been killed by the fumes, most of the survivors were males. These were 
resting or flying about near the openings when the cisterns were examined. On 
several occasions in January I was bitten by females disturbed from cisterns. 


Examinations were continued in April in parts of Haifa not previously visited. 
Larvae were found to be present in nearly every case, together with adults. Usually 
some larvae were obtained in the first bucket of water drawn. After that it was 


MOSQUITOS COLLECTED IN PALESTINE. 391 


difficult to secure many more, even though in one case twenty buckets of water were 
examined in succession. When disturbed they evidently move away from the 
vicinity of the opening. As far as I could judge, from a number of observations 
made, the larvae of A. bifurcatus living in these situations are chiefly surface feeders, 
existing upon debris which may be washed in by the rains. 


I successfully reared adults from the egg by keeping the larvae in a bowl of clean 
water and feeding them upon a diet of chopped flies. A few young larvae found 
on 18th November fed up slowly in the same way, living for ebout two months 
before pupating. The adults prefer cool and dark places in which to rest during 
the day, and were not found in any abundance in houses or tents. 


Except for occasional individuals the species is absent from Egypt. Specimens. 
were obtained in the following localities from sea-level to about 3,000 feet :-— 


PALESTINE : Jerusalem; Bethlehem; Jaffa; Haifa; Acre; Nazareth. Syria: 
Baalbek ; Zahle ; Beirut. 


Anopheles algeriensis, Theo. 

I think this must be the species referred to by previous workers in Palestine as 
A, fragilis, Theo., or A. aitkent, James. It does not appear to be very general, and I 
have no records for Egypt or Cilicia. 


One female was captured on 3rd December 1919 in a tent pitched in the marshes 
east of Acre; this had recently sucked blood. On 11th and 12th August 1920 it 
was found in numbers at Wadi Selhab, a marshy tract about three miles south of 
Jenin on the Nablus road and over twenty miles from the coast. The females 
were biting freely at sunset, and some fifty specimens were caught in half an hour. 
The only other mosquitos seen at the same time were A. hyrcanus and Culex pipiens 
(one). 

It would appear to breed chiefly in the larger marshes, away from human habita- 
tions, like A. hyrcanus, and is therefore probably not a frequent carrier of malaria. 
It has been found in the marshy areas of the Auja valley, between Mulebbis and 
Jaffa. 


A few specimens were bred from larvae found in a small marsh on the outskirts of 
Beirut, in September 1919. 


Anopheles hyrcanus, Pall. (sinensis, Wied. ; pseudopictus, Grassi). 

As mentioned above, this species breeds chiefly in the larger marshes and is to be 
found along the coastal belt of Palestine, at least as far north as Acre. It also occurs 
in suitable localities inland. I think there is no doubt that it is to be found in Syria 
and northwards, but I did not meet with it. I believe that up to the present it has’ 
not been found in Egypt. 

The adults may be found nearly all the year round, but in larger numbers in winter 
and spring. 

PALEsTINE: Marshes in the Auja Valley, near Mulebbis ; Athlit ; the Kishon 
Plain, from Haifa to Acre ; Wadi Selhab. near Jenin. 


392 CAPT. P. J. BARRAUD. 


Anopheles mauritianus, Grandpré. 

This is one of the Anopheles which is to be found regularly in Palestine and Feypt. 
In the former country it breeds in much the same environment as A. hyrcanus, viz. 
the swamps of the Auja and Kishon rivers. I have not met with it inland, or in 
Syria. In Egypt it is found chiefly in the lake margins and swamps of the Delta. 


The adults appear to be most common in the early part of the year. 


Anopheles superpictus, Grassi (palestinensis, Theo. ; nurse, Theo.). 

Generally distributed over Palestine and Syria, and met with in Cilicia. The 
larvae are usually found in natural water, and prefer clear pools, near springs of 
fresh: water. 

The adults are no doubt responsible for a great deal of the prevailing malaria in 
country districts, as distinct from towns. My collection includes specimens from 
the following localities, from 1,100 feet below sea-level to about 3,000 feet above :— 

PALESTINE: Sarona (near Jafla); Ludd, Latron, Ram Alla (near Jerusalem) ; 
along the course of the river Auja from Rasel Ain to Mulebbis ; Beisan ; Tabgha ; 
at many places along the Jordan from Beirut Yakub bridge, on the Upper Jordan, 
down to Ghoranyieh (near the Dead Sea). Syria: Ak river, near Marash ; Zahle, 
Lebanon Mountains. Criicta: Bozanti; Toprak Kali, Taurus Mountains. 


Anopheles culicifacies, Giles, var. sergenti, Theo. 

This mosquito was not discovered until the end of my stay, and I think it is 
probably often overlooked amongst numbers of the preceding. A few females were 
found in tents at Tabgha, on 15th August 1920. Previously a few specimens were 
bred from larvae found in pools along the Nazareth road, near Haifa. 


Anopheles multicolor, Camb. (chaudoyei, Theo.). 

This insect has been previously known to workers in Palestine as A. turkhudi, 
Liston. It is now found to be distinct from the Indian species. 

The larvae are often found in large numbers in brackish marshes along the coastal 
plain of Palestine, in late autumn, spring and early summer. It is not, however, 
confined to the coast, and occurs in salty pools and streams in the Jordan Valley. 
As mentioned on a previous page, it is considered to be the chief malaria-carrier in 
Egypt, where it has a wide distribution. 


Anopheles pharoensis, Theo. 

Although abundant in some parts of Egypt, it is regarded as a doubtful carrier 
of malaria. In October 1919 I found it commonly at Kantara, on both sides of the 
Canal, and also in the northern suburbs of Cairo. It is very rare in Palestine, and 
T obtained only one female. This was found at Tabgha, north of Tiberias, on 15th 
August 1920. I have no records for Syria. 


Stegomyia fasciata, F. 
This is exclusively a domestic species, found in most parts of Egypt, Palestine 
and Syria, at least as far north and east as Aleppo. 


— ve 


MOSQUITOS COLLECTED IN PALESTINE. 393 


Larvae were found in water receptacles of all descriptions, including flower-vases 
in a sitting-room, fire-buckets containing clean water, and a disused boiler in a 
kitchen. . 

The adults are most troublesome in the hot season, and are suspected of being 
carriers of the organism causing “Aleppo sores” and “Jericho boils.” Some 
of the Palestine specimens are very beautiful, having the dorsal surface of the 


abdomen almost completely covered with silvery scales. 


PALESTINE: Jaffa; Jerusalem; Haifa; Jenin; Tiberias. Syria: Beirut ; 
Damascus ; Aleppo. 


Ochlerotatus caspius, Pall. (dorsalis, Theo., nec Mg.) 

A very abundant and widely distributed species in Egypt, but so far as my 
observations go, it is not so common in Palestine. 

The larvae prefer brackish water. 

A few adults were found at Haifa in June 1919, 

The night of 31st March 1920 was spent at Jisr 2l Damie, Jordan Valley. During 
the evening we were attacked by clouds of mosquitos, but owing to an unfortunate 
accident to my supply of tubes, I was unable to bring back a large number of speci- 
mens. A few of this species were secured, together with O. detritus, Theobaldva 


annulata var., and Anopheles superpictus. 


Ochlerotatus mariae, Serg. 

One of the few mosquitos to be found breeding in sea-water. Large numbers 
of ova, larvae and pupae were found in pools above high water mark on the rocks 
at Athlit, Palestine, in August 1920. In June and July 1919 larvae were abundant 
in similar situations along the coast to the south-west of Beirut, Syria. The pools 
become very salt owing to evaporation, and the temperature of the water, by 
exposure to the sun, is usually well over 90° F. for the greater part of the day. 


The adults bite freely in the daytime, and cause much annoyance to people bathing 
from the shore. 


Ochlerotatus detritus, Hal. (salinus, Fic.). 
A very abundant and troublesome species in parts of Palestine during the wet 
season and spring. 


The larvae are found chiefly in brackish water, near the coast, but I also found 
them in large numbers in the Jordan valley at the end of March 1920, at 1,000 feet 
below sea-level. 


PALESTINE: Zummarin; Haifa; Acre; Jisr el Damie, Jordan. 


Culex pipiens, L. 
Abundant and widely distributed in Egypt, Palestine and Syria, and found 
as a rule near human dwellings. The species occurs from sea-level to 4,500 feet. 


394 CAPT. P. J. BARRAUD. 


The larvae differ in some respects from Western European and Mesopotamian 
specimens, and appear to represent a distinct race. No diflerences can, however, 
be discovered in the adults. Larvae collected in Jerusalem, Haifa and Baalbek 
(Syria), are very uniform in structure and size and show the following obvious 
differences from typical specimens :—Full-grown larva smaller, with distinctly 
shorter siphon ; slightly smaller average number of pecten teeth ; antennae light, 
instead of dark. In one case larvae were found living in a well in which the water- 
level was 85 feet below the ground. 

The adults exhibit a certain amount of variation in the abdominal banding, in 
some specimens the white bands being incomplete dorsally. 


[Culex fatigans, Wied. | 

Although this mosquito was recorded from Ezypt by several of the earlier workers, 
it does not seem to be found there. An examination of the genitalia of all males 
in the collection of the Egyptian Ministry of Agriculture has proved them to be 
C. pipiens, I have been unable to find the species in Palestine or Syria. 


Culex laticinctus, Edw. 

The larvae may sometimes be found in very large numbers in covered surface 
wells, tanks, and similar situations in Palestine. 

The adults have been observed in quantities in tents in camps. In Egypt it is 
found commonly at Alexandria. 

PALESTINE: Jerusalem ; Wadi Hamish; Haifa. Syria : Baalbek. 


Culex univittatus, Theo. 

Distributed from Egypt to Syria, but never m any great abundance. The 
larvae live in marshes and stream pools, generally away from dwellings. One 
is lable to overlook them, as, when disturbed, they immediately leave the surface, 
returning for only a second or two at long intervals. 

There still seems to be some confusion regarding the larval characteristics. My 
specimens disagree in almost every particular with the description and figure given 
in Bull. Ent. Res. x, p. 69, 1919; neither do they fit in with the synoptic table 
given in the same journal vol. i, p. 381, Dec. 1912. I have handed some of my 
material to Mr. F. W. Edwards, who has kindly offered to go into the matter, and. 
publish the results in a future paper. 

PALESTINE: Wadi Hamish; Sarona; Haifa. Syria: Beirut. 


Culex tipuliformis, Theo. 

Found in similar situations to the preceding. It has not, so far as | am aware, 
been recorded from Egypt under this name, but there is a specimen from the Fayum. 
in the British Museum collection. 

PALESTINE : El Afule; Acre; Haifa. Syria : Damascus. 


MOSQUITOS COLLECTED IN PALESTINE. 395 


Culex hortensis, Fic. 

Although I was on the look out for this species in Palestine, very few were found. 
Some females were taken at Haifa in May 1920. No doubt it is common enough 
in some parts of the country. At Damascus, in July 1919, larvae of this and of 
C. tupuliformis were dipped from the same pool. It appears to be absent from 


Egypt. 


Culex mimeticus, Noé. 
Not uncommon in suitable localities in Palestine, Syria and Cilicia, but absent 
from Egypt. The larvae have been found associated with those of A. swperpictus. 


PALESTINE: Wadi Hamish. Syria: Beirut. CrLicta: Bozanti. 


(Culex modestus, ['ic. | 
This was not met with, but is likely to occur, since it has been found in Macedonia 
and Egypt. 


Theobaldia longiareolata, Macq. 

One of the commonest gnats in Egypt, Palestine and Syria. It is so universal 
that it is hardly necessary to give localities. This and the following species were 
found up to 4,500 feet in the Lebanon Mountains. 

The larvae abound in water-barrels and other receptacles in gardens, surface 
wells, and covered and uncovered tanks and cisterns. 


Theobaldia annulata, Schrank. 

Much less common than the last-named, and absent from Egypt. 

The strongly marked type form was taken at Ramleh, Palestine, and at Ain 
Sofar, Lebanon Mountains. In the Jordan Valley at 1,000 feet below sea-level, 
I found some specimens of a pale variety which agree with the Mesopotamian 
form, referred to in my paper on mosquitos of that region (Bull. Ent. Res. x, p. 325, 
1920). 


Theobaldia morsitans, Theo. 
At the end of April 1920 some larvae were found in a pool at the bottom of a 
small quarry at Haifa. The species was not met with elsewhere. 


[Taeniorhynchus richiardii, Fic. | 
This species has been found in Palestine, at Jell Zahmfil, Huleh, by Dr. J. Cropper, 
but was not met with by the author. 


Uranotaenia unguiculata, Edw. 

Widely distributed in Egypt, and found in parts of Palestine. The larvae as a 
tule are only to be found in the larger swamps. 

My thanks are again due to Dr. Guy A. K. Marshall, Director of the Imperial 
Bureau of Entomology, and Mr. F. W. Edwards, of the British Museum, for assistance 
in the identification and synonymy of certain of the species and forms mentioned 
above. Also to Mr. G. Storey, of the Egyptian Ministry of Agriculture, for informa- 
tion as to the distribution of mosquitos in Egypt. 


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397 


A PRELIMINARY REVISION OF THE GENUS DOCIOSTAURUS, FIEB. 
By B. P. Uvarov, 


Assistant Entomologist, Imperial Bureau of Entomology. 


The genus Dociostaurus, Fieb., which is synonymous with Stawronotus, Fisch., 
includes several species of locusts and grasshoppers injurious to agriculture in South- 
Kastern Europe, Central and Western Asia and North Africa, the well known 
Moroccan locust (Dociostawrus maroccanus, Thunb.) being one of the worst pests 
in Algeria, Tunisia, Asia Minor, the Caucasus and Turkestan. The systematics of the 
species of this genus are in a very unsatisfactory state, and this, together with the 
tendency of the species to individual variability, is the cause of many mistakes in 
their identification on the part of economic entomologists. The object of this paper 
is, therefore, to establish a more or less natural system of the species enabling every- 
one to identify them with certainty. 

The materials upon which this work is based were assembled by me from different 
sources. First ofall, I have worked out the collection of the Tiflis Museum (Caucasus), 
where very large series of Dociostaurus species from Transcaucasia and Persia ‘are 
to be found. Then also the very rich collection of the Turkestan Entomological 
Station (in Tashkent) was placed at my disposal by the Director of this Station, 
V. Plotnikov. My friend N. Ikonnikov sent me large series of specimens from 
Turkestan, Bokhara, and other places, as well as the material in the Museum 
of Moscow University. The Entomologist of the Entomological Station in Bairam- 
Ali (Transcaspia), D. A. Smirnov, collected for me in that locality, where I myself 
had also collected previously. My own collection contains extensive series of this 
genus taken by myself in Uralsk province (S. HE. Russia), Northern Caucasus and 
Transcaucasia. Very helpful for me was the kindness of Messrs. C. & I. Bolivar, 
-who sent me for study several Spanish and North African representatives of this 
genus, as well as the types of species described by I. Bolivar. To all the above- 
named gentlemen I wish to express here my cordial thanks for the loan of 
material which has enabled me to accomplish this revision. 


The genus Dociostaurus was described in June 1853 by Fieber (Lotos, 1, p. 118), 
and in the same year, but in November, by Fischer (Orthoptera EKuropaea, pp. 297, 
351) under the name Stauronotus ; Fischer’s name has been generally accepted, 
but it must give way to Dociostaurus, Fieb., according to the law of priority. 
W. F. Kirby in his catalogue (Synon. Cat. of Orthoptera, i, pp. 152-154, 1910) 
includes in the genus Dociostawrus 12 species. Later on, I. Bolivar described in 
1911 D. kerviller from Persia and in 1917 D. dantini from Morocco; Kirby and 
Waterhouse included (Fauna Brit. India, Acrid., pp. 117-120, 1914) in this genus 
five species described by Walker under Stenobothrus. Prof. Stchelkanovzev 
described in 1909 D. tartarus (Izvestia Varshav. Universit., Warsaw, 1909, p. 34), 
which is omitted in Kirby’s Catalogue. Lastly, F. Werner published in 1913 one 
more species from Egypt under the name of D. gracilis (Zool. Jahrb., Syst., xxxiv, 
p- 220). 

(737) F2 


398 B. P. UVAROV. 


Thus the number of species of the genus Dociostaurus reached 19. This number, 
however, must be reduced to 10 only, since some of the species do not really belong 
to this genus and others again must be regarded as synonymous, as follows :— 

Oedipoda hastrio, Fisch. Wald. (Orth. Imperii Rossici, p. 311, pl. 32, fig. 6) belongs, 
according to Fischer Waldheim, to the section of his genus “ Oedipoda”’ with three 
keels on the pronotum, the two lateral keels being straight or nearly so; this is 
not the case in the genus Dociostaurus, one of its chief generic characters being the 
strongly angulated side keels of the pronotum. I am of opinion that Fischer 
Waldheim’s species is merely one of the numerous colour forms of Calliptamus 
atalocus, L. 

All five of Walker’s species, as a study of the types in British Museum clearly 
shows, belong to the genera Avzlacobothrus, Bol. and Leva, Bol., and not to 

_Dociostaurus.* 

_ Stauronotus australis, Bol., can hardly belong to this genus, since it is described 
from tropical West Africa, while all representatives of the genus are palaearctic 
in their distribution. 

Stauronotus hauensteom, Bol., and Stauronotus kerviller, Bol., are synonymous 
with Dociostaurus crassiusculus, Pantel, as I am convinced from a study of Bolivar’s 
types. 

Stauronotus tartarus, Stchelk., is not an independent species, but a geographical 
race of Dociostaurus crucigerus, Ramb., as is shown below. 

Docwostaurus gracilis, Werner, though very unsatisfactorily described, is most 
likely to be synonymous with D. genez, Ocsk. 

Unfortunately but few species of this genus (for example, maroccanus) are sharply 
characterised, while the rest of them are extremely variable individually, their 
variability being expressed not in coloration or dimensions only, but to some extent 
in morphological characters as well. In consequence of this, it 1s quite easy to make 
mistakes in studying representatives of such extremely variable species as crassius- 
culus or anatolicus, unless very large series of specimens are available. Though I 
have been able to study rather long series (more than one thousand specimens 
altogether), I still think it impossible to give yet a full monograph of the genus, and 
this revision must be regarded as but a preliminary one. 


* The synonymy of these species of Walker’s is as follows :-— 
» Leva apicalis (Walker). ‘ 
1871. Stenobothrus apicalis, Walker, Cat. Derm. Salt. Brit: Mus., v, Suppl., p. 80. 
~ 1871. Stenobothrus turbatus, Walker, l.c. p. 81. 
1910. Dociostaurus apicalis, Kirby, Syn. Cat. Orth., i, p. 154, no. 11. 
1910. Dociostaurus turbatus, Kirby, l.c., p. 154, no. 12. 
1914. Dociostaurus apicalis, Kirby, Fauna Brit. India, Acrid., p. 117, no. 135, fig. 90. 
1914. Dociostaurus turbatus, Kirby, l.c., p. 118, no. 136. 
Leva mundus (Walker). 
1871. Stenobothrus mundus, Walker, Cat. Derm. Salt. Brit. Mus., v, Suppl., p. 79. 
~ 1871. Stenobothrus epacromoides, Walker, l.c., p. 81. 
1914. Dociostawrus mundus, Kirby, Fauna Brit. India, Acrid., p: 119, mo: 137. 
1914. Dociostaurus epacromoides, Kirby, l.c., p. 119, no. 138. 
2.1918. Stawroderus exemplaris, J. Bolivar, Rev. R. Acad. Se. Madrid, xvi, p. 380. 
Aulacobothrus decisus (Walker). 
1871. Stenobothrus decisus, Walker, Cat. Derm. Salt. Brit. Mus., v, Suppl., p. 80. 
1914. Dociostaurus decisus, Kirby, Fauna Brit. India, Acrid., p. 120, no. 139, fig. 91. 


A PRELIMINARY REVISION OF THE GENUS DOCIOSTAURUS. 399 


The genus Dociostaurus is purely palaearctic in its distribution, and none of the 
species is to be found beyond the borders of the Palaearctic Region. The centre 
of development of the genus is to be looked for in the dry table-lands of Western 
and Central Asia, where the largest number of species occurs and where they are 
especially disposed to individual variability. From there diffetent species spread 
(and are spreading continually) in all directions, but more especially into the 
plains of the vast palaearctic desert belt, extending through Northern Africa, and 
South-western and Central Asia, in the fauna of which species of Dociostaurus 
present one of the most characteristic features. The distribution of each species 
will be discussed below in more detail. 


Key to the Species. 

1(10). Hind femora attenuate, narrow, slightly incrassate at the base. Pronotum 
constricted in the middle; white stripes not dilated in metazona, or 
altogether absent. 

2 (3). Elytra extending considerably beyond the apex of abdomen. Antennae 
in both sexes but slightly extending beyond the hind margin of pronotum. 
Fore tibiae thickened. (Foveolae of the vertex narrowed anteriorly. Front 
vertical ; its ridge above ocellum sulcate in male and plane in female. 
Hind femora with three very distinct black spots outwardly and black 
knees. Hind tibiae sanguineous, with base pale—Length of body, $ 20-28, 
2 28-38 ; of elytra, J 20-27, 9 25-36 ; of hind femora, ¢ 15, 2 20 mm.) 

1. maroccanus, (Thunb.) 

3 (2). Elytra not or scarcely reaching the apex of hind femora. Antennae of male 
one and half times or even twice as long as the head and pronotum taken 
together ; antennae of female extending considerably beyond the hind 
margin of pronotum. Fore femora not thickened. 


4 (5). Head not at all or but slightly prominent above pronotum. Foveolae of the 
vertex longer than wide. Front in male slightly inclined, with flat ridge, 
in female vertical with ridge convex above ocellum. Elytra reaching the 
apex of hind femora. Hind femora with three sharp black spots 

2. crucigerus (Ramb._). 

a(d). Hind tibiae red. Black marks very distinct. 
b(c). General habitus slender ; headnotthick. (Length of body, 3 14-16, 
© 18-238 ; of elytra, g 9-11, 9 15-17; of hind femora, $10, 2 13 mm.) 
2a. crucigerus crucigerus (Ramb.). 
c(b). General habitus more clumsy ; head thick. (Length of body g 12-16, 
© 20-24 ; of elytra § 8-10, 7 12°5-15 ; of hind femora ¢ 10:2, 9 12-15 
mm.) ; 2 .. 2b. crucigerus hispanicus (Bol.). 
d(a). Hind tibiae as, ‘Cotoesion more or less pale, with the black and 
white markings indistinct or totally absent. (Dimensions as in crucigerus) 
2c. crucigerus tartarus (Stchelk.). 

5 (4). Head very prominent above, especially in male. Foveolae of the vertex 
as long as wide. Front in male strongly, in female markedly reclinate. 
Elytra not or scarcely reaching the apex of hind femora. 


400 


6 (7). 


7 (6). 
8 (9). 


9 (8). 


13(12). 


B. P.. UVABOV 


Occiput without carina in its middle. Hind tibiae bluish. Foveolae of 
the vertex not narrowed anteriorly. (Antennae in both sexes one and half 
times as long as head and pronotum together. General coloration pale 
greyish, with grey markings ; hind femora with rather distinct grey spots.— 
Length of body, $12, 215-17; of pronotum, $ 2-3, 93; of elytra, 
§ 10-12, 9 13-14; of hind femora, § 6-7, 99-11 mm.) .. 3. gener (Ocsk.). 

Occiput with a short but distinct longitudinal carina in the middle. 


Small, slender. Foveolae of the vertex strongly narrowed anteriorly. 
Frontal ridge in male impressed along its whole length, in female quite 
flat. Hind tibiae yellowish or bluish, without dark ring near the base. 
(General coloration brownish grey or clay-coloured. Antennae in male 
twice, in femate one and half times as long as head and pronotum together. 
Wings hyaline or bluish.—Length of body, gj 12-14, 2 13-16; of elytra, 
¢ 7-10, 9 11-13; of hind femora, 3 8-10, 9 11-13 mm.) 

4, albicornis (Eversm.). 


Larger and heavier built. Foveolae of the vertex not or but slightly 
narrowed anteriorly. Frontal ridge in male quite flat, im female convex 
above ocellum, but flat and disappearing below it. Hind tibiae greyish, 
bluish or rose, with a black or dark ring near the base. (General coloration 
very variable. Antennae in male twice as long as in female but a little 
longer than head and pronotum together. Wings rosy or bluish—Length 
of body, ¢ 18-27, 9 23-32 ; of elytra, J 12-16, 2 15-20; of hind femora, 
3 12-15, 9 15-20mm.).. 7 s Ly 5. anatolicus (Krauss). 

Hind femora short, wide, rather incrassate at the base. Pronotum cylindri- 
cal, not constricted in the middle; white stripes in metazona dilated. 


Rather slender, with head prominent above. Hind tibiae never red. 


. Foveolae of the vertex longer than wide, not narrowed anteriorly, and 


contiguous at the apex. Frontal ridge in male plane, with slight impression 
below ocellum (female unknown). (Black markings on pronotum and 
hind femora sharp. Hind tibiae yellowish —Length of body, g 12°5; 
of elytra, $5°5; of hind femora, ¢9mm.) .. .. 6. dantini, Bol. 


Foveolae of the vertex nearly as long as wide, narrowed anteriorly. Frontal 
ridge in male sulcate for its whole length, in female flat and slightly im- 
pressed near ocellum. (General coloration clay-like, with very indistinct 
castaneous markings. Hind tibiae bluish—Length of body, 914, 221; 
of elytra, ¢ 5-8, 9 8-15; of hind femora, ¢ 10, 9 14 mm.) 

7. plotnikovi, sp. n. 

Clumsy, with thick head, not prominent above. Hind tibiae red or 
sanguineous, 


Lateral carinae of pronotum present not only on metazona, but also on 
fore part of prozona. d: anal segment with two large rounded contiguous 
lobes behind, covering the base of the supra-anal plate; the latter rounded, 
with apex attenuate. ©: anal segment far wider than long; supra-anal 
plate triangular, its length equal to its basal width. Hind tibiae coral-red. 


16(17). 


17(16). 


18(15). 


A PRELIMINARY REVISION OF THE GENUS DOCIOSTAURUS. 401 


Smaller. Foveolae of the vertex distinctly impressed, markedly narrowed 
anteriorly. Front rather reclinate; frontal ridge in profile straight. 
(Yellowish-grey or clay-coloured, with more or less distinct grey and pale 
markings ; lateral lobes of pronotum with a black square spot in the middle. 
Antennae in male one-half, in female slightly, longer than head and pro- 
notum together. Elytra not longer than abdomen, but very often far 
shorter.—Length of body, § 12-18, 9 18-26; of elytra, g 7-11, 29-18; 
of hind femora, ¢ 9-10, 912-15mm.).. 8. crassvusculus (Pantel). 

Larger. Foveolae of the vertex but slightly impressed, nearly square. 
Front nearly vertical; frontal ridge in profile convex. (Straw-yellow, 
with sharp grey and black markings. Antennae in female as long as head 
and pronotum together, in male a little longer. Elytra reaching the middle 
of abdomen.—Length of body ¢ 23, 2 32; of nai $8, 213; of hind 
femora, g 13, 218 mm.) a 2 9. kurdus, sp. n. 

Lateral carinae of the pronotum aentat on metazona only. 4: anal 
segment with two small, widely separated lobes behind ; supra-anal 
plate triangular, not attenuate at the apex. Q: anal segment as long as 
wide ; supra-anal plate longer than its basal width. Hind tibiae sanguin- 
eous. (Clayish grey or brown, dull, with indistinct grey and pale markings. 
Lateral lobes of pronotum with a small square black spot m the middle. 
Elytra reaching the apex of abdomen.—Length of body, 3 16-18, 9 23-25 ; 
of elytra, g 11-12, 9 14-16; of hind tibiae, J 11, 9 15 mm.) 

10. raussi (Ingen.). 


Annotated Catalogue of Species. 


1. Dociostaurus maroccanus (‘Thunb.). 


1815. 
1824. 
1825. 
1832. 
1836. 


1846. 


1853. 
1853. 
1854. 
1873. 
1882. 


1910. 


Gryllus maroccanus, Thunberg, Mem. Acad. Petersb., v, p. 244. 

Gryllus maroccanus, Thunberg, l.c., ix, pp. 399, 425, no. 91. 

Gryllus crucvatus, Charpentier, Horae Entom., p. 137. 

Oedipoda cruciata, Brullé, Exped. Morée, Ins., p. 92, no. 63, pl. 30, fig. 2. 


Acrydium cruciatum, Costa, Fauna Regni Napoli, Acrid., p. 37, no. 17, 
pio, far 1; A, yc. 


Oedipoda vastator, Fischer Waldheim, Orth. Imp. Ross., p. 312, no. 30, pl. 14, 
fig. 1. 

Docwostaurus cruciata, Fieber, L.dfon! i, p. 118 (partem). 

Stauronotus cruciatus, Fischer, Orth. Europ., p. 352 (partim), pl. 17, fig. 11, a, b. 

Dociostaurus cruciata, Fieber, Syn. Kur. Orth., p. 18, no. 1 (partim). 

Stauronotus maroccanus, Stal, Rec. Orth., p. 111, no. 1. 


Stauronotus maroccanus, Brunner v. Wattenwyl, Prodr. Europ. Orth., p. 136, 
Ho: ©. 


Dociostaurus maroccanus, Kirby, Syn. Cat. Orth., ui, p. 152, no. 1. 


This species was described by Thunberg in 1815 and 1824, and redescribed twice 
since—by Charpentier as Gryllus cruciatus* and by Fischer Waldheim as Oedipoda 


*1I am obliged to Dr. W. Ramme, who kindly studied for me Charpentier’s type of 
cruciatus in the Berlin Museum and informed me that it is identical. with maroccanus, 


Thunb. 


402 - B. P. UVAROV. 


vastator. Fieber and Fischer evidently described this species (under cruciatus) 
together with crucigerus of Rambur, which is a quite distinct species; and the 
same mistake has been repeated even by Brunner, who says that maroccanus is a 
very variable species, which is not the case. In fact, I had the opportunity of study- 
ing many hundreds of specimens from different localities and may say that the 
individual or geographical variability in this species 1s extremely small, and it is very 
constant in all its characters. It can very easily be recognised by the aid of the key 
given above. From its nearest relative, D. crucigerus, it may be distinguished by its long 
elytra, short antennae and thickened (in males) fore femora, besides its different. 
dimensions. It is interesting to mention that females during the period of oviposition 
gradually change their coloration, which becomes dull clay-coloured, the black 
markings turning to light brown. This coloration might be regarded as protective 
while the female is sitting on the ground, but it is more likely to depend on physio- 
logical causes. The coloration of the males changes in this period as well, but to 
a far less degree. 

Geographical Distribution. The range of distribution of the Moroccan locust 
includes North Africa, Southern Europe, Syria, Palestine, Arabia, Mesopotamia, 
Asia Minor, Armenia, Persia and the Aralo-Caspian plains. Within this vast region 
there are to be found several places where this species finds most favourable natural 
conditions and where it becomes a serious pest of agriculture, breeding in large 
masses and acquiring swarming habits.* 


2. Dociostaurus crucigerus (Ramb.). 

1838. Gryllus crucigerus, Rambur, Faune Entom. d’Andal., ui, p. 86, no. 13. 

1848. Oedipoda brevicollis, Eversmann, Addit. quaedam laevia ad Fisch. Wald. 
celeb. Orth. Ross., p. 11, tab. A, fig. 4. 

1853. Dociostaurus cruciata, Fieber, Lotos, 1, p. 118 (partim). 

1853. Stauronotus cruciatus, Fischer, Orth. Europ., p. 352 (parti). 

1853. Stauronotus crucigerus, Fischer, l.c., p. 356, no. 5. 

1854. Dociostaurus cruciata, Fieber, Syn. Europ. Orth, p. 18, no. 1 (partim). 

1859. Stauronotus cruciatus, Kversmann (nec Charp.), Bull. Soc. Nat. Moscow, 
a (Pp. AST. 

1860. Stauronotus genei, Tiirk (nec Ocsk.), Wien. Ent. Mon., iv, p. 86, no. 3. 

1862. Stauronotus annulipes, Tiirk, l.c. vi, p. 81. : 

1867. Stauronotus brevicollis, Frivaldsky, Ertek Termész. KGr., 1 (12), p. 162, no. 1, 
pl 6, fis."10, 

1882. Stauronotus brevicollis, Brunner v. Wattenwyll, Prodromus Europ. Orth., 
pp. 136, 137, no. 2, pl. 5, fig. 30, A, B. 

1910. Docrostaurus brevicollis, Kirby, Syn. Cat. Orth., ii, p. 153, no. 4. 

Rambur’s description of Gryllus crucigerus leaves no doubt that he had before 

him not D. maroccanus, Thunb. (known at that time as cruciatus, Charp.), but 

another species distinguished by its smaller size, shorter elytra and with the lateral’ 

carinae of the pronotum present on the prozona as well as on the metazona—1.e.. 


* The biology of D. maroccanus is fully described by W. La Baume in a ari 4 
published book. 


A PRELIMINARY REVISION OF THE GENUS DOCIOSTAURUS. 403 


the species described ten years later by Eversmann as Oedipoda brevicollis. After- 
wards, however, Fieber and Fischer, who evidently did not know Rambur’s species, 
confused it, as well as brevicollis, Ev., with cruciatus, Charp. (maroccanus, Thunb.). 

This miscomprehension of crucigerus has been accepted by all following writers ; 
even Eversmann himself in 1859 regarded his brevicollis as synonymous with 
cruciatus, Charp. Later on brevicollis, Ev., was raised to specific rank by Frivaldsky 
(1867), but the identity of brevicollis and crucigerus, Ramb., has never been 
recognised. Rambur’s name, however, has ten years priority over brevicollis and 
must therefore be adopted. The same species has been again described in 1860 
by Tiirk under Stawronotus genet (nec Ocsk.), and two years later the author changed 
this name, as preoccupied by Ocskay, into annulipes. 

This species in its typical form is easily recognisable from its nearest relative— 
maroccanus—by the characters given above, which are very constant. 

Geographical distribution. The area of distribution of this species occupies the 
southern (Mediterranean) parts of Europe, South Russia, Southern Siberia, Turkes- 
tan, Transcaspia, the Caucasus, Asia Minor, Persia and North Africa. It is to be 
mentioned, however, that the species is to be sub-divided into three different 
geographical races (subspecies), as follows :— 


2a. Dociostaurus crucigerus hispanicus, Bol. 
1898. Stauronotus brevicollis var. hispanicus, I. Bolivar, Ann. Sci. Nat. Porto, 
¥;.{pe, 14. 

This subspecies, peculiar to Spain, is distinguished from the typical form by its 
more heavy habitus and especially its thick head (thanks to the kindness of Messrs. 
Bolivar I have had the opportunity of studying this form). It would be of interest 
to examine a large series of specimens from Southern France, whence Rambur’s 
type came. If specimens from this locality are identical with hispanicus, then 
the Eastern European form, which I take now for the typical, must have another 
name, and brevicollis, Kv., might be applied to it as a subspecific name. 


2b. Dociostaurus crucigerus crucigerus, Ramb. 

The typical form occurs over the whole of Southern Europe, South Russia, 
Caucasus (except the Eastern plains of Ciscaucasia and Transcaucasia) and Southern 
Siberia. | 


2c. Dociostaurus crucigerus tartarus, Stschelkan. 

1909. Stauronotus tartarus (Sauss. in litt.), Stschelkanovzev, Izvestia Imper. 
Varshav. Univers., Warsaw, 1909, p. 34, no. 22. 

——. Stauronotus gener, auctorum (nec Ocsk.), partim. 

This south-eastern desert subspecies is easily recognised by the colour of its hind 
tibiae, as well as the rather indistinct and extremely variable general coloration. 
On account of its pale tibiae it is very often confused with D. genei, Ocsk., which 
has good morphological differences from it. The area of distribution of this race 
lies in desert plains of the Aralo-Caspian basin, Persia, Mesopotamia and Northern 
Africa ; in fact it populates the whole Eremian subregion of the Palaearctic Region. 


404 B. P. UVAROV. 


The economic importance of Dociostawrus crucigerus, with its subspecies, is 
not .so great as that of maroccanus, since it is a non-swarming and non-migratory 
species, but still it is one of the injurious grasshoppers and does some damage to 
grass and corn crops in Spain and Western Siberia, together with different species 
of Stenobothrus and other grasshoppers. 


3. Dociostaurus genei (Ocsk.). 
1832. Gryllus gener, Ocskay, Acta Acad. Leop.’Carol, xvi (2), p. 961. 


1853. Acridium pygmaeum (Bonelli in litt.), Fischer, Orth. Europ., pp. 352, 353, 
tab. xvii, fig. 11c, 11d. 

1910. Dociostaurus gener, Kirby, Syn. Cat. Orth. mi, p. 154, no. 8. 

21913. Docrostaurus gracilis, Werner, Zoolog. Jahrb., Syst., 34, p. 220. 

Closely related to albicorns, Ev., but always recognisable by the lack of the 
longitudinal carina on the vertex. From D. crucigerus tartarus it may be distinguished 
by the prominence of the upper part of the head. 

Geographical Distribution. This is not quite clear, since the species has been often 
confused with albicornis and crucigerus tartarus, but it seems that it 1s a native 
of the desert plains of Western Asia and North Africa, occurring also in some isolated 
places in Southern Europe. 


There is nothing known about the economic importance of this species. 


4. Dociostaurus albicornis (Eversm.). 

1848. Stauronotus albicorns, Eversmann, Addit. quaedam laevia ad Fisch. Wald. 
Orth. Ross., p. 10, no. 6, pl. A, fig. 3. 

1910. Dociostaurus albicornis, Kirby, Syn. Cat. Orth., ui, p. 157, no. 9. 

Though the difference between albicornis and genet is rather considerable, it 
seems desirable that an examination should be made of large series of specimens 
of both species, which may then prove to be conspecific. 

Geographical Distribution. Not so extensive as that of genet, since it is known 
only from the Aralo-Caspian basin; its occurrence in Germany, recorded by Azam 
(Bull. Soc. Ent. France, 1913, p. 218) is almost certainly based on a wrong deter- 
mination or a mistake in date. 

The economic importance of this species is unknown. 


5. Dociostaurus anatolicus (Krauss). 

1896. Stauronotus anatolicus, Krauss, Zoolog. Jahrb., Syst., ix, p. 560, no. 7, pl. 8, 
fio. 1, 

1896. Stauronatus anatolicus var. castaneopicta, Krauss, l.c., p. 562, no. 7a, pl. 8, 
fig. 2, 2 A, B. 

1907. Stauronotus anatolicus var. castaneopicta, Adelung, Horae Soc. Entom. 
Ross., xxxviii, p. 51, no. 10, pl. 1, fig. 2. 

1910. Dociostaurus anatolicus, Kirby, Syn. Cat. Orth., itl, p. 153, no. 7. 


A PRELIMINARY REVISION OF THE GENUS DOCIOSTAURUS. 405 


An extremely variable species in its size, relative dimensions and coloration. The 
form described and figured by Krauss and Adelung under the name v. castaneopicta 
is remarkable for the coloration of the head and especially of the pronotum and 
elytra, on which large velvet-castaneous spots are to be seen on a straw-yellow 
ground. 

Geographical Distribution. D. anatolicus occurs in Asia Minor, Eastern and 
Southern Transcaucasia, the whole Aralo-Caspian basin and Persia. 

In some localities of Turkestan it occurs in large quantities, though it does not 
form swarms, and may cause some damage to grass; no attacks on cultivated 
crops, however, have been recorded. 


6. Dociostaurus dantini, I. Bol. 
1914. Dociostaurus dantini, I. Bolivar, Mem. R. Soc. Espafi. Hist. Natur., vi 
(5a), p. 186. 
A very distinct species, as I have been able to ascertain by a study of the unique 
type specimen. 
Geographical Distribution. Described from Morocco. 


7. Dociostaurus plotnikovi, sp. nov. 

The short diagnosis of this new species given above may be completed by the 
following description :— 

Form slender. General coloration clay-yellow, with obsolete grey markings. 
Head elongate, prominent above (especially in male) ; eyes elongate ; front in male 
strongly reclinate and wholly sulcate, in female markedly reclinate, flat, with a slight 
impression near the ocellum. Antennae in male one and half times as long as head 
and pronotum together, in female only a little longer. Foveolae of the vertex markedly 
impressed, nearly as long as wide, narrowed in front. Typical sulcus of pronotum 
placed behind the middle; hind margin of pronotum rotundate ; disk obsolete 
castaneous (very often only in metazona) with pale lateral cross-shaped lines, which 
are triangularly widened on the metazona ; lateral lobes with an obliterated grey 
or castaneous spot in the middle. Elytra about as long as abdomen, unicolorous or 
with scarcely distinct grey spots. Hind femora with typical spots greyish, 
obsolescent. Hind tibiae bluish or pale. ¢@: anal segment with two small rotundate 
widely separated lobes; supra-anal plate rotundato-triangular; subgenital plate 
short, with blunt apex, very hairy. Q: anal segment wider than long; supra-anal 
plate rotundato-triangular. (Dimensions, see Key.) 

Geographical Distribution. This species has been sent to me from the Turkestan 
Entomological Station by the Director, V. Plotnikov, from Turkestan (Golodnaya 
Steppe) and Bokhara (Mirshade-Karatag), where it occurs sometimes in considerable 
quantities, and its larvae even form migrating swarms, causing damage to different 
cultivated crops. 


8. Dociostaurus crassiusculus (Pant.). 


1886. Stauronotus crassvusculus, Pantel, Ann. Soc. Esp. Hist. Nat., xv, p. 237, pl. 2, 
fig. 1. 


406 B. P. UVAROV. 


1893. Stauronotus hauensteini, Bolivar, Rev. Biol. Nord. France, v, p. 480. 

1894. Stauronotus hauensteinr, Giglio-Tos, Boll. Mus. Torino, ix (191), p. 2. 

1896. Stauronotus hauensteim, Krauss, Zoolog. Jahrb., System., ix, p. 562, no. 9, 
pl. 8, fig. 3. 

1898. Stauronotus crassiusculus, Bolivar, Ann. Sci. Nat. Porto, v, p. 13,14. 

1902. Stauronotus crassiusculus, Navas, Bol. Soc. Esp. Hist. Natur., ii, p. 335, no. 3. 

1910. Dociostaurus hauensteini, Kirby, Syn. Cat. Orth., ii, p. 153, no. 2. 

1910. Dociostaurus crassiusculus, Kirby, l.c., p. 153, no. 6. 

1911. Dociostaurus kervillet, Bolivar, Bull. Soc. Sci. Nat. Rouen, xlvii, p. 36. 

1913. Stauronotus crassiusculus, Pant., var. cappadocicus, Azam, Bull. Soc. Ent. 
Frances 1913p. 219: 

D. crassiusculus, Pantel, was described from Spain, and hauensteini, Bol., from 
Asia Minor, but I have most carefully studied topotypic specimens of both species 
and could not find any difference between them. I had very large series of this 
species from Asia Minor, Transcaucasia and Persia, which enable me to state that. 
it is very variable in its dimensions and coloration, though not in morphological 
characters, which are very stable. D. kervillei, described by Bolivar more recently, 
is but a small and rather light-coloured form of the same species, as | am convinced. 
from a study of the type specimen. The variety described by Azam is a mere 
individual aberration which is not worth a distinctive name. 


Geographical Distribution. This is as yet not fully known. It seems that this 
species has two separate areas of distribution : one in Spain and another in Western 
_ Asia (Asia Minor, Armenia, Eastern Transcaucasia, Persia, Syria); but it is quite 
possible that it occurs in North Africa as well. If even these two areas are actually 
separated, there is nothing unnatural in such distribution of a species, since we know 
that there existed once a closer connection between the eastern and western parts 
of the Mediterranean countries than is now the case. 

There are no records concerning the economic importance of this species. 


9. Dociostaurus kurdus, sp. nov. 

There is nothing to be added to the short diagnosis of this species given above, 
since it agrees in all essential characters with D. crassiusculus, the differences being 
given in the key to the species. These differences are, in fact, so few that I should 
have been inclined to treat kurdus as only a geographical form of crassvusculus were it 
not that it has also some important morphological features, and I prefer to regard it 
in the meantime as a distinct species. 

Geographical Distribution. I had a number of specimens of both sexes from Muséék 
and Vezne, in Kurdistan, taken by P. Nesterov in June and July, 1914. 


10. Dociostaurus kraussi (Ingen.). 


1897. Stauronotus kraussi, Ingenitsky, Horae Soc. Entom. Ross, xxxi, p. 63, pl. 7. 
1910. Dociostaurus krausst, Kirby, Syn. Cat. Orth., ii, p. 153, no. 3. 


A PRELIMINARY REVISION OF THE GENUS DOCIOSTAURUS. 407 


This species is very well characterised by numerous morphological features, and 
is quite easy to recognise. It is not very variable in dimensions and little more so 
in coloration, but all the morphological characters are very constant. 

Geographical Distribution. D. kraussvis distributed all over the Kirghiz Steppes 
(north from the Caspian Sea), Turkestan, Transcaspia, Hastern Ciscaucasia and South- 
western Siberia. In the latter country it is one of the commonest grasshoppers, 
being sometimes very injurious to pastures and corn crops. 


A supplement to the synonymy of Dociostaurus maroccanus, Thunb. 

In the course of rearranging the collection of Locustidae in the British Museum, 
I came across the type of Epacromia oceanica, Walker (Cat. Derm. Salt. Brit. 
Mus., wv, p. 779, No. 26, 1870), which has been included by Kirby (Syn. Cat. 
Orth., ui, p. 193, No. 12) in the genus Chortoicetes, but is undoubtedly con- 
specific with D. maroccanus. The type specimen of FH. oceanica bears the label 
“ Sandw. I., Beechey ”; but there are in the British Museum a few more un- 
doubtedly palaearctic species bearing the same label, and the only explanation is 
that these specimens have been taken during Capt. Beechey’s journey from 
Europe, via the Canaries, to the Sandwich Islands, most probably at the Canaries 
or at Madeira, and that they have been wrongly labelled. 


J 
i i. i" 
rth 

cane 


409 


COLLECTIONS RECEIVED. 

The following collections were received by the Imperial Bureau of Entomology 
between Ist October and 3lst December 1920, and the thanks of the Managing 
Committee are tendered to the contributors for their kind assistance :— 

Mr. T. J. Anderson, Government Entomologist :—36 Culicidae, 199 Tabanidae, 
96 Stomoxys, 242 other Diptera, 795 Coleoptera, 214 Hymenoptera, 2 Planipennia, 
112 Lepidoptera, 10 species of Coccidae, 235 other Rhynchota, 41 Orthoptera, 
12 Odonata, and 47 Ticks ; from Kenya Colony. 

Mr. E. Ballard, Government Entomologist :—8 Microlepidoptera ; from Madras. 

Mr. Cecil N. Barker, Durban Museum :—29 Bruchidae and 2 Hispidae ; from 
South Africa. 

Capt. P. J. Barraud, Entomologist to the Egyptian Expeditionary Force :—1 
Dorylus Ant and 5 Odonata ; from Palestine. 

Dr. Chas. K. Brain :—2 Diptera ; from Stellenbosch, South Africa. 

Division of Entomology, Pretoria :—6 Diptera, 52 Coleoptera, 8 Hymenoptera, 
and 49 Rhynchota ; from South Africa. 

Mr. P. R. Dupont, Curator of the Botanic Station :—2 species of Coccidae ; from 
Seychelles. 

Mr. T. Bainbrigge Fletcher, Imperial Entomologist :—150 Bees ; from India. 

Mr. C. C. Gowdey, Government Entomologist :—7 Diptera, 59 Coleoptera, 135 
Ants, 33 other Hymenoptera, 5 Lepidoptera, 10 Thrips, 38 species of Coccidae, 
490 Aphididae, 7 other Rhynchota, 80 Isoptera, 5 Orthoptera, 5 Odonata, 7 Ticks, 
and 1 Scorpion ; from Jamaica... 

Mr. G. F. Hill, Entomologist, Australian Institute of Tropical Medicine :—4 
Isoptera ; from North Queensland. 

Mr. M. Afzal Husain, Government Entomologist :—57 Jassid Bugs and 2 
Orthoptera ; from the Punjab. 

Mr. J. C. Hutson, Government Entomologist :—323 Ticks ; from Ceylon. 

Dr. A. Ingram :—97 Culicidae, 100 Psychodidae, 8 Auchmeromyia, 9 Cordylobia, 
180 other Diptera, 50 Siphonaptera, 3 Coleoptera, 2 Planipennia, 11 Lepidoptera, 
and 30 Mites ; from the Gold Coast. 

Mr. G. W. Jeffery :—15 Lepidoptera ; from Lumbwa, Kenya Colony. 

Dr. W. A. Lamborn :—4 Cordylobia, 64 Chalcids, and 21 Coleoptera; from 
Tanganyika Territory: and 8 microscope-preparations of Stegomyia scutellaris 
infested with protozoal parasites, 52 other preparations, 19 Coleoptera, and 5 
Orthoptera ; from the Federated Malay States. 

Major W. F. M. Loughnan, R.A.M.C. :—A number of blood-sucking Chironomidae 
from Jamaica. 

Dr. J. W. Scott Macfie :—Preparations of Toxorhynchites mosquitos ; from Accra, 
Gold Coast. 

Mr. G. Mer, Jewish Health Bureau :—4 Culicidae and 2 Tabanidae:; from Palestine. 

Mr. Norman C. E. Miller :—2 species of Coccidae ; from Morogoro, Tanganyika 
Territory. 


410 COLLECTIONS RECEIVED. 


Mr. W. H. Patterson :—1,589 Coleoptera and 17 Lepidoptera; from the Gold 
Coast. : 

Mrs. W. H. Patterson :—46 Diptera, 1,735 Coleoptera, 29 Lepidoptera, and 
25 Rhynchota ; from the Gold Coast. 

Mr. A. H. Ritchie :—463 Coleoptera ; from Jamaica. 

Mr. W. Robson, Curator of the Botanic Station :—Specimens of the leaf blister 
mite of cotton ; from Montserrat. 

Senhor A. F. de Seabra :—3 Tabanidae, 4 other Diptera, 101 Coleoptera, 25 
Hymenoptera, and 28 Rhynchota ; from San Thomé. 

Mr. F. R. Shepherd, Agricultural Superintendent :—Specimens of the leaf blister 
mite of cotton ; from St. Kitts. 

Mr. Hubert W. Simmonds :—47 Diptera, 2 Hymenoptera, 5 Micro-lepidoptera, 
and 4 Rhynchota ; from Fiji. 

Mr. 8. H. Skaife :—59 Bruchid beetles and 12 Chalcid parasites ; from South 
Africa. ; 

Mr. O. H. Swezey, Entomologist to the Hawaiian Sugar Planters’ Association :— 
12 Weevils ; from Hawai. 

Me. F. W. Urich :—365 Coleoptera, 2 Hymenoptera, 6 species of Coccidae, and 20 
Aleurodidae ; from San Thomé. 

Mr. R. Veitch :—10 Coleoptera, 8 Hymenoptera, 21 Lepidoptera, and 11 
Rhynchota ; from Fiji. 

Wellcome Bureau of Scientific Research :—6 Coleoptera ; from Peru. 


ae 


- 


=— 


411 


GENERAL INDEX. 


abdominalis, Coccinella. 


A butilon, food-plant of thrips in Sudan, 
98. 


Acacia spp., in 8.E. Africa, 320. 


Acanthus montanus, Tabanids associ- 
ated with flowers of, in Katanga, 147. 


Acartomyia (Aédes) zammittii, breed- 
ing in salt water in Dardanelles, 261. 


Accra, breeding places of mosquitos in, 
258, 259, 261, 265. 


acinosa, Lachnosterna. 

Aclerda, 2, 15. 

Aclerda digitata, on grass in Natal, 15. 

aconitus, Anopheles (Myzomyia) mini- 
Mus. 


Adina microcephala, in S.E. Africa, 
320, 376. 


Aédes argenteus (see Stegomyia fasciata). 


»  (Skusea) cartroni, in Madagascar, 
134. 


» fluviatilis, breeding in salt water 
in Panama, 261. 


»  (Ochlerotatus) fryeri, 134. 

». mnigeriensis (see Ochlerotatus). 
»  (Skusea) pembacnsis, 134. 

»  punetothoracis (see Ochlerotatus). 
» eammiti (see Acartomyia). 


Aédomyia africana, in West Africa, 74, 
269. 


Aédomyia squamipennis, in Panama, 
73. 


Aegophagamyia terticeps, in Aldabra, 43. 
aegyptiaca, Lcerya. 
Aesculus hippocastanwm (Horse Chest- 


nut), mosquitos breeding in, in 
Italy, 259. 

aethiopica, Corizoneura  (Tabanus, 
Tanyglossa). 


(741) 


Africa, New Aphids from, 65-72; 
new lLonchaeidae from, 199-210; 
classification of Tabanidae of, 139— 
152. 


Africa, East, new weevils from, 274. 


Africa, Portuguese East, bionomics of 
tsetse flies in, 315~—385. 


Africa, South, Coccidae of, 1-41; new 
Aphids from, 65-72; Tabanidae 
from, 141, 143, 152. 


Africa, West, bionomics of mosquitos 
in, 74, 105-112, 247-270. 

africana, Aédomyia; Filippia (see 
F. carissae) ; Stegomyia. 

africanum, Lecanium. 

africanus, Ceroplastes (see C. mimosae) ;. 
Mansonioides; Phlebotomus minutus- 

agricola, Tanymecus. 

aitkeni, Anopheles (Stethomyia). 

albicornis, Dociostaurus. 

albifacies, Buplex (Corizoneura). 

albimanus, Anopheles. 

albipectus, Tabanus. 

Albizzia chirindensis, in S.E. Africa, 
321, 322, 382. 

Albizzia lebbek, mosquito larvae breed- 
ing in, in Lagos, 257. 

albopicta, Stegomyia. 

albotaeniatus, Anopheles 
chus). 

Aldabra, Tabanids from, 45. 

Algeria, Myelois phoenicis in, 288 ; 
bionomies of Parlatoria blanchardt 
in, 296, 297. 

algeriensis, Anopheles. 

alienum, Lecanium (see L. hesperidum). 

Allopulvinaria, gen. n., 2, 16. 

Allopulvinaria subterranea, on grass in 
Cape Colony, 16. 


(Myzorhyn- 


B 


-412 


-alluaudi, Bouvierella (Scione). 
-alpinus, Ochlerotatus. 
amanda, Ocnerogyia. 


amaryllidis, Lecanium assimile (see 
L. hesperidum). 


America, Central, Lachnosterna setifera 
in, 193. 


Amirante Is., Tabanus albipectus from, 
45. 


Amomum, in S.E. Africa, 322. 
-ampelinus, Rhizoctonus. 
ampelorhiza, Schizoneura. 


Anacardium occidentale (Cashew), mos- 
quito larvae breeding in, in Lagos, 
257. 


anatolicus, Dociostaurus. 


angustatum, Lecanium (see L. hesperi- 
dum). 

angustipes, Simulium. 

angustitarsis, Simulium. 

annetti, Taeniorhynchus. 

annulata, Theobaldia ; Uranotaenia. 

annulioris, Culex. 

annulipes, Stauronotus. 

annulitarsis, Culex. 

anodentata, Lachnosterna. 


Anopheles, in Lagos, 247-270; in 
Malay Archipelago, 77-92 ; breeding 
among water lettuce in Panama, 
73-75. 


Anopheles aconitus (see A. minimus). 


a (Stethomyia) aitkeni, distri- 
bution of, in Malay Archi- 
pelago, 91; A. algeriensis 
recorded as, in Palestine, 
391. 


us aitkent var. insulae-florum, 
larva of, in Java, 81. 


. aitkeni var. papuae, in New 
Guinea, 81. 


# albimanus, breeding among 
water-lettuce in Panama, 
74. 


2 (Myzorhynchus) albotaenia- 
tus, larva of, in Sumatra 
and Java, 82, 83. 


i algeriensis, in Palestine, 
391. 
as (Nyssorhynchus) annulipes 


var. moluccensis, in 
Moluccas and NewGuinea, 
78, 79, 90; larva of, 80. 


GENERAL INDEX. 


Anopheles barberi, breeding in trees 
in America, 259. 


os (Myzorhynchus)  barbiros- 


tris, larva of, 86. 


ss (Myzorhynchus) barbiros- 
tris var. pallidus, larva 
of, in Sumatra and Java, 
82. 


. barianensis (see A. plum- 
bews). 


a bifurcatus, in Palestine and 
Syria, 368, 390. 


chaudoyet (see A. multi- 
color). 


. costalis, breeding places of: 
in West Africa, 74, 251— 
270. 


“7 culicifacies var. sergenti, in 
Palestine, 392. 


ee (Stethomyia) _culiciformis, 
81; breeding places of in 
India, 259. 

9 fragilis (see A. algeriensis). 

sf (Nyssorhynchus) fuliginosus, 
larva of, 87. 

- (Nyssorhynchus) fuliginosus 
var. nivipes, characters 
of, 88. 

“ funestus, breeding places 
of, in Lagos, 260, 269, 
270; nm Madagascar, 133. 


: hyrcanus, in Palestine, 391, 
392; larva of, 389. 

7 (Myzomyia) immaculatus, in 
Java, ii: 

” ms indefinitus (see 
A. vagus). 

S3 (Cellia) kocht, 85. 

9 (Neomyzomyia) leucosphy- 


rus, larva of, in Malay 
Archipelago, 83, 84. 

a (Myzomyia) ludlowi, breed- 
ing in salt water in 
Andamans, 261; dis- 
tribution of, in Malay 
Archipelago, 89, 91. 

maculipalpis, in Mauritius, 
133. 

maculipennis, in Palestine, 
388-390; in North Russia, 
198. 

marshalli, breeding places 
of, in West Africa, 74, 
259; in Madagascar, 133. 


39 


GENERAL INDEX. 413 


Anopheles mauritianus, breeding 


“99 


99 


% 


9? 


99 


9? 


99 


places of, in West Africa, 
262-270; in Mauritius, 
133; larvaof, in Palestine, 
389, 392. 

(Myzomyia) minimus, larva 
of, 88. 

minimus var. aconitus, in 
Malay Archipelago, 85, 
86, 88, 89. 

minimus var. cohaesa, 89. 


multicolor, breeding in salt 
water in Egypt, 261; in 
Palestine, 388, 392; 
larva of, 389. 

nigripes (see A. plumbeus). 

nili, breeding places of, in 
West Africa, 267—270. 

nurse (see A. superpictus). 

obscuratus, in West Africa, 
269, 270. 


palestinensis (see A. super- 
pictus). 

(Myzorhynchus) peditaenia- 
tus, 82. 


pharoensis, in Lagos, 269 ; 
in Madagascar, 133; in 
Palestine and Egypt, 392. 


plumbeus, distribution of, 
259. 

pseudopictus (see A. hyr- 
canus). 

(Nyssorhynchus) punctula- 


tus var. tesselatus, larva 
of, 85, 89, 90. 


rossi (see A. subpictus). 


(Nyssorhynchus) schiffneri, 
in Malay Archipelago, 
87, 88, 90. 

(Myzorhynchus) sinensis, in 
Malay Archipelago, 82, 
86, 87, 91; (see A. hyr- 
canus). 


squamosus, in Madagascar, 


(Myzomyia) subpictus (ros- 
st), breeding in salt water 
in India, 261; in Malay 
Archipelago, 89, 90. 

superpictus, in Palestine, 
388, 389, 392, 393. 

(Cellia) tananariviensis (see 
A. squamosus). 

tarsimaculatus, breeding 
among water-lettuce in 
Panama, 74. 


Anopheles transvaalensis, in Madagas- 
ear, 133. 


re (Myzorhynchus) wmbrosus, 
in Malay Archipelago, 84. 

a (Myzomyia) vagus (indefini- 
tus), in Malay Archipel- 
ago, 87, 89. 


Anoplodesmus saussurei, an imported 
pest in Mauritius, 175. 


antennatus, Phlebotomus. 


anthurui, Chermes (see Saissetia hemis- 
phaerica). 


antigonm, Pulvinaria. 

Antigua, new Lachnosterna from, 191. 
antiguae, Lachnosterna. 

anxifer, Culex (see CO. fatigans). 
Aonidia blanchardi (see Parlatoria). 
aonidum, Chrysomphalus. 

Apalis, in §.E. Africa, 368. 


Aphelinus mytilaspidis, parasite of 
Parlatoria blanchardi, 296, 297. 


Aphididae, new African, 65-72; from 
Persia, 153. 


Aphis buxtoni, sp. n., in Persia, 153. 


»  punicae, on pomegranate in 
Persia, 155, 156. 


»  punicellae, on Punica granatum 
in Egypt, 156. 

»  repariae, on Vitis, 66. 

»  sorghi, in Sudan, 95. 

»  vitis, on Vitis, 66. 
apicalis, Leva (Stenobothrus). 
apicoannulatus, Ochlerotatus. 
apicoargentea, Stegomyta. 


appendiculata, Pangonia (see Cori- 
zoneura aethiopica). 


arboricollis, Orthopodomyia 
Newsteadina). 


arcticum, Simulium. 
arcuatus, Culicoides. 


Arenipses sabella, distribution of, in 
stored dates, 289, 290. 


argenteopunctatus, Culex (Heptaphle- 
bomyia) ; Ochlerotatus. 


argenteostriatum, Simulium. 

argenteus, Aédes (see Stegomyia fascsata). 

argyreatum, Simulium. 

Aristolochia, Coccids on, in Mauritius, 
174. 

aristolochiae, Pulvinaria. 

Arizona, date pests in, 288, 296. 


(Culex, 


B2 


414 


GENERAL INDEX. 


artemisiae, Lecanium (Coccus) (see | balfouri, Uranotaenia. 


Ceroplastes rusct). 
articulatus, Aspidiotus. 


Artocarpus incisa (Bread-fruit), 
mosquito larva breeding in, in Lagos, 
257 ; Coccids on, in Mauritius, 174. 

arundinis, Hyalopterus. 


Arvicanthis, in Glossina areas in S.E. 
Africa, 338. 


Asia, new Lonchaeidae from, 199-210. 
asparagi, Lichtensia. 


Aspidiotus articulatus, on Phoenix in 
Africa, 299. 


- betulae, parasitised by 
Aphelinus mytilaspidis in 
Italy, 297. 

+ cyanophylli, 
299. 


on Phoenix, 


$3 destructor, on guavas in 
Mauritius, 173. 

a dictyospermi, on Phoenix, 
299. 

-: jicus, on Phoenix, 299. 

~ hederae, on date palm in 
California, 299. 

lataniae, on Phoenix, 299. 

- mauritianus, on palms and 
Cycas in Mauritius, 173. 

- ostreaeformis, on date palm 
in California, 299. 

a rapax, on Phoenix. 299. 


assimile, Lecanium (see L. hesperidum). 


Asterolecanium spectabile, on palms and 
Cycas in Mauritius, 173. 


ataeniatus, Culex. 
ater, Isaniris. 
atratula, Lonchaea (Carpolonchaea). 


Aulacobothrus decisus, synonomy of, 
398. 


aurantii, Chrysomphalus. 

aurea, Lonchaea (Lamprolonchaea). 
aureosimile, Simulium. 

aureum, Simulium. 

aureus, Taeniorhynchus. 

aurites, T'aeniorhynchus. 

austen, Glossina; Simulium. 
Australia, Lonchaea in, 204-207. 
australis, Stawronotus. 

Avocado Pear (see Persea gratissima). 


Bamboos, Coccids on, in Mauritius, 
hy &e8 


bancrofti, Filaria. 


Banksinella lineatopennis, in Lagos, 
269. 


” punctocostalis, 
268, 269. 


barbata, Osca (Pangonia). 
barberi, Anopheles. 
barbirostris, Anopheles. 


barianensis, Anopheles (see A. plum- 
beus). 


Bauhinia spp., in 8.E. Africa, 320, 322,. 
326, 349. 


beaumontiae, Lecanium (see Saissetia. 
hemisphaerica). 


beckeri, Pangonia. 

begoniae, Lecanium 
nigra). 

betranus, Systates. 

benefactrix, Mutilla. 

Bersama niassae, in S. E. Africa, 348 - 

betulae, Aspidiotus. 

bifurcatus, Anopheles. 

bipartitus, Ceroplastes. 

biroi, Lonchaea. 

bisignatus, Tabanus tropicus. 

bisulcata, Lonchaea (Carpolonchaea). 

bituberculatus, Ceroplastes. 


blanchardi, Parlatoria 
Websterella). 


Blighia sapida (Akee Apple), mosquito- 
larvae breeding in, in Lagos, 257. 

boisduvali, Diaspis. 

Bombylius, 151. 

bougainvilleae, Cerciaphis. 

Bouvierella allwaudi, in Seychelles, 43. 
4 inornata, sp. 0., m. Sey- 

chelles, 44. 
borealis, Tabanus. 
botulibranchium, Simulium. 


Brachylaena rhodesiana, in 8. E. Africa, 
322, 348. 


Brachystegia, 124; new weevil on, 
in Rhodesia, 273; influence of, on dis- 
tribution of Glossina in 8. E Africa, 
320, 321, S225 A2eawee,, aos G2, 
348, 850; .351,.379: 


bracteatum, Simulium. 


in Lagos, 


(see Sarssetia. 


(Aonidia,. 


GENERAL INDEX, 


Bradyornis ater, enemy of Glossina in 
S.E. Africa, 368. 


” murinus, enemy of Glossina 
in §.E. Africa, 368. 


Brahmina, considered a synonym of 
Lachnosterna, 189. 


brassicae, Pulvinaria (see P. floccifera). 


Brazil, breeding places of mosquitos in, 
254. 


Bread-fruit (see Artocarpus). 


brevicollis, Oedipoda (see Dociostaurus 
crucigerus). 


brevipalpis, Glossina ; Toxorhynchites. 
Bridelia micrantha, in 8.E. Africa, 320. 


Britain, bionomics and classification 
of Stmuliwm spp. in, 211-246. 


brouniana, Lonchaea (see DD. awrea). 
brunnipennis, Buplex (Pangonia). 


Bucorax cafer, a potential host of 
Glossina in 8.E. Africa, 328. 


Buphaga, enemy of eeene in S8.E. 
Africa, 369. 


Buplex, gen. n., 139; African Tabanids 
placed in, 140. 


»  juscinervis, sp. N., 
Colony, 140. 


a (Pangonia) suavis, 141. 


in Cape 


Burkea africana, relation of,” to 
Glossina morsitans in §.E. Africa, 
365. 


buxtoni, Aphis. 


cacti, Chilocorus. 

Cadicera, a synonym of Phara, 140. 
caecutiens, Chrysops. 

Calandra glandium, sp. n., in acorns in 


India, 277. 

rs sculpturata, in South Africa, 
277. 

- shoreae, sp. n., food-plants 
of, in India and Mauritius, 
276. 


caleitrans, Stomoxys. 

calidella, E'phestia. 

California, Coccid pests of date palms 
in, 

caliginosus, Ochlerotatus. 

Callinectes marginatus, 248. 

Calliptamus italicus, 398. 


Calophyllum inophyllum, mosquito 
larva breeding in, in Lagos, 257. 


415 


Calymnatus hesperidum (see Lecanium),. 
Calypticus hesperidum (see Lecanium). 


_ hydatis (see Ceroplastes 
TUSCt). 

a laevis (see Lecanium 
hesperidum). 

E radiatus (see COeceroplastes 
TUsct). 

a testudineus (see Ceroplastes 
TUSCL). 


candela, Ceroplastes. 
cantator, Culex. 


Capitophorus chrysanthemi, on Chrysan- 
themum in Orange River Colony, 69. 


caraciae, Chermes (see Ceroplastes rusct). 


Cardisoma armatum (Land Crab), 
mosquitos breeding in 
holes of, in Lagos, 247. 


A guanhumi, C. armatum 
possibly identical with, 
247. 


carduellinum, Rhopalosiphum. 

Carica papaya (Pawpaw), mosquito 
larva breeding in, in Lagos, 257. 

caricae, Coccus (see Ceroplastes rusct). 

carissae, Filuppia. 

carpenteri, Pangonia. 

Carpolonchaea, sub-gen. n., 199. 

Carpophilus dimidiatus, in dates, 294. 


cartroni, Aédes (Skusea, Stegomyia) ; 
Culex (see C. fatigans). 


carueli, Diaspis. 

Caspian coast, Aphids from, 154, 155. 
caspius, Ochlerotatus. 

cassinae, Lecanium (see Saissetia oleae) | 


caucasicus, Phlebotomus (see P. 
gent). 


cautella, E'phestia. 
Cavariella, 154. 


Cephalophus monticola, in 8.E. Africa, 
327, 338. 


Ceratopogoninae, breeding in crab- 
holes in Brazil, 254. 


Ceratophyllum, preferred by larvae of 
Simulium, 230. 


Cerciaphis, gen. n., 65, 70. 
af bougainvilleae, Sp. D., On 
Bougainvillea in Uganda, 70. 


Cercopithecus spp., relation of, to 
distribution of Glossina in S.E. 
Africa, 336, 349. 


Sere 


416 


certferus, Ceroplastes (see C. destructor). 


Cerodonta dorsalis, seasonal variation 
of 214: 


Ceronema, 2, 22. 


- mobilis, sp. n., on Celastrus 
cordata in Natal, 22. 


Ceroplastes, 2, 26. 


- africanus (see C. mimosae). 

ss bipartitus, 26. 

- candela, in Natal, 27. 

= ceriferus (see C. destructor). 

= combreti, sp. n., on Com- 
bretum in Transvaal, 27. 

5: destructor, on custard 
apple, etc.,in Transvaal, 
28. 

* egbarum, on Acacia in 


Natal, 28. 
fulleri, in Natal, 
29. 


99 39 


ay eucleae, Sp. n., in Trans- 
vaal, 30. 

“ longicorda, sp. n., in Natal, 
ol: 

-- mimosae, on Acacia 
karroo in Cape Colony, 
31. 

‘3 myricae, on Myrica quer- 


cifoia in Cape Colony, 
32. 


>» pallidus, sp. n., on fig in 
Transvaal, 33. 
> quadrilineatus var. sim- 


plex, n., on Rhus in Cape 
Colony, 33. 


>> rusci, On quince in Cape 
Colony, 34. 
i tachardiaformis, sp. D., on 


Elytropappus rhinocero- 
tis in Cape Colony, 35. 


= zonatus, on Acacia in 
Transvaal, 35. 


Ceroplastodes, 2, 40. 


a bituberculatus, sp. n., in 
Cape Colony, 40. 


Ceylon, Lonchaea in, 209; Aphelinus 
mytilaspidis parasitisng Parlatoria 
blanchardi in, 297 ; Icerya aegyptiaca 
infesting dates in, 299. 


chaudoyei, Anopheles (see A. multi- 
color). 


GENERAL INDEX. 


Chermes anthurii (see Saissetia hemis- 


phaerica). 
caraciae (see Ceroplastes rusct).. 
+ hesperidum (see Lecanium). 
- laurt (see Lecanium hesperi- 
dum). 
nd oleae (see Saissetia). 


chilianthi, Filippia. 

Chilocorus cacti, predaceous on Parla- 
toria blanchardi in America, 297. 
Chionaspis graminis, parasitised by 
Aphelinus mytilaspidis 

in Ceylon, 297. 


“ permutans, parasitised by 
Aphelinus mytilaspidis 
in Ceylon, 297. 

x pinifolii, parasitised by 
Aphelinus mytilaspidis 
in U.S.A., 297. 


“A simplex, on bamboo in 
Mauritius, 173. 


chirindensis, Systates. 
chrysanthemi, Capitophorus. 


Chrysomphalus aonidum, on date palm. 
in California, 299. 


‘5 aurantvi, on date palm. 
in California, 299. 


Chrysops, in North Russia, 196, 197. 


2 caecutiens, in North Russia,. 
197, 198. 

3 divaricata in North Russia, 
198. 

3 nigripes, in North Russia, 
198. 

Pe relicta, in North Russia, 198. 

ds sepulcralis, in North Russia, 
198. 


Chrysophyllum flavum, relation of 
Glossina to, in 8.E. Africa, 319. 


chrysostigma, Phara (Pangonia). 
Cilicia, mosquitos in, 390, 392, 395. 


Cinnamomum zeylanicum, Coccids on,. 
in Mauritius, 174. 


cisst, Macrosiphum. 
Cissus, new Aphids on, 66. 
cttricola, Lonchaea (Carpolonchaea). 


Citrus aurantium (Orange), mosquito: 
larvae breeding in, in Lagos, 257. 
clypeatum, Lecanium (see Sarssetia 

hemisphaerica). 
Coccidae, South African, 1-41. 


GENERAL INDEX. 


Coccinella abdominalis, predaceous on 
Parlatoria blanchardi in America, 297. 


Cocos nucifera (Coconut), mosquito 
larvae breeding in, in Lagos, 257. 


Coccotrypes dactyliperda, in dates in 
Egypt, 294. 


Coccus artemisiae (see Ceroplastes rusci). 
»  caricae (see Ceroplastes rusct). 


»  coffeae (see Saissetia hemis- 
phaerica). 


»  cycadis (see Saissetia oleae). 
»  floccifera (see Pulvinaria). 

»  hesperidum (see Lecanium). 
»  myricae (see Ceroplastes). 

» oleae (see Saissetia). 

»  palmae (see Saissetia oleae). 


»  patelliformis (see Lecanium 
hesperidum). 


»  rusci (see Ceroplastes). 
cockerelli, Pheidole. 


coffeae, Lecanium (Coccus) (see Saissetia 
hemisphaerica). 


cohaesa, Anopheles 


minimus. 
columbaczense, Simulium. 


Columnea myricae (see Ceroplastes). 


: testudinata (see Ceroplastes 
TUSCt). 
- testudiniformis (see Cero- 


plastes rusci). 
combreti, Ceroplastes. 


Combretum, in relation to distribution 
of Glossina in S.E. Africa, 320, 345, 
349; Coccid on, in Transvaal, 27. 


cometes, Lachnosterna (Phytalus). 


. condei, Hretmopodites (see EH. quinque- 
vittatus). 


consimilis, Culex. 
continentalis, Lonchaea. 
Conofilippia, gen. n., 2, 25. 


subterranea, sp. 0., in 
Transvaal, 25. 


in relation to 
im toch. 


Conopharyngia spp., 
distribution of Glossina 
Africa, 319, 348. 


Corchorus olitorius, food-plant of thrips 
in Sudan, 98. 


Cordia interrupta, food-plant of Tiphia 
parallela in Mauritius, 164. 


(Myzomyia) 


417 


Corizoneura, systematic position of, 
1 


s aethiopica, Synonymy of, 
139. 
_ albifacies (see Buplezx). 
ie dissimilis (see Buplex). 
7 distincta, 140. 
formosa, sp. n., in Cape 
Colony, 142. 
33 (Diatomineura)  hastata, 
140. 
” % amornata, 
140,145, 
146,147.. 
9 (Pangonia) lateralis, 140. 
ns (Diatomineura)  lineati- 
thorax, 140. 
35 (Pangonia) longirostris, 
140. 
a (Diatomineura) neavet, 
140. 
Ks pallidipennis, 140. 
99 (Diatomineura) pene- 
trabilis, 140. 
» (Pangonia) sagittaria, 140 
9 (Pangonia) taprobanes, 
140. 
s schwetzi, sp. n., in Central 
Africa, 143, 146, 147. 
a umbratipennis, 140. 
99 (Diatomineura) virgata, 
140. 


corniger, Culex. 
costalis, Anopheles. 
Cotton Thrips (see Heliothrips indicus). 


Crabs, mosquitos breeding in holes of, 
in Lagos, 247, 248. 

crassicornis, Haematopota. 

crassipalpis, Phara (Pangonia). 

crassiusculus, Dociostaurus. 

in S.E. 


Craterispermum  laurinum, 


Africa, 320. 
Crateropus, in 8.E. Africa, 365, 368. 


s kirki, possibly destroying 
Glossina pupae in §.E. 
Africa, 369. 


Cremastogaster scutellaris, associated 
with Aphids in Persia, 156. 


Crinum giganteum, mosquito larvae 
breeding in, in Lagos, 257. 


crocodilinus, Tabanus. 


418 


Crotalaria, 


GENERAL INDEX, 


food-plant of thrips in 


Sudan, 98. 
erucigerus, Dociostaurus. 
Oryptinglisia, 2, 38. 


lounsburyi, on Vitis in 
Cape Colony, 38. 


Oucumis melo, food-plant of thrips in 
Sudan, 98. 


Culex, breeding among water lettuce, 


3° 


39 


3° 


3° 


39 


39 


3° 


- he) 


3) 


39 


9° 


3°) 


39 


3° 


3) 


3) 


>> 


74; larvae of, attacked by 
Inspa, 279-281. 


annulioris, 136. 

annulitarsis, 136. 

anxifer (see C. fatigans). 
arboricollis (see Orthopodomyia). 


argenteopunctatus, in Mada- 


gascar, 137. 


ataeniatus, breeding places of, 
in Lagos, 264, 265, 269. 


(Aédes) cantator, breeding in 
salt water in America, 261. 
cartront (see C. fatigans). 


corniger, breeding in crab-holes 
in Brazil, 254. 

consimilis, breeding places of, 
in Lagos, 253-269. 


decens, 137; early stages of, in 
West Africa, 105, 251-269; 
pupa of, 106, 107, 108. 


duttoni, breeding places of, in 
Lagos, 259-269. 


extricator, in Tropical America, 
254. 


fatigans, in Madagascar and 
Mauritius, 137; breeding 
places of, in Lagos and Gold 
Coast, 253-269; probably 
not occurring in Egypt, 394. 


fluviatilis (see Aédes). 
flavus (see Taeniorhynchus). 


fowleri (see Ochlerotatus nigeri- 
ensis). 


giganteus, in Madagascar, 136. 


grahami, breeding places of, in 
Lagos, 259-269. 


grandidieri (see Taeniorhynchus 
flavus). 


guiarti, breeding places of, in 
Lagos, 264, 265, 269. 

hortensis, in Palestine, 395. 

insatiabilis (see Stegomyia 
fasciata). 


Culex insignis, breeding places of, in 


99 


Lagos, 251-253, 264-269. 
invidiosus (see C. decens). 
laticinctus, in Palestine and 

Egypt, 394. 
laurenti, in Madagascar, 137. 
lividocostalis (see C. decens). 
mimeticus, in Palestine and 

Syria, 395. 
modestus, probably occurring in 

Palestine, 395. 
neireta (see C. giganteus). 
nigrocostalis (see C. decens). 
pipiens, in Madagascar, 137 ; 

in Palestine, 391, 393, 394. 
pruina, in Lagos, 268, 269. 
quasigelidus, in West Africa, 

74, 268, 269 ; in Madagascar, 

136. 
reticulatus, in Brazil, 254. 
rima, breeding places of, in 

Lagos, 251-254, 268, 269. 
ronaldi (see C. sittens). 
salinarius, breeding in salt 

water in America, 261. 


salisburiensis, breeding places 
of, in Lagos, 251-253, 268, 
269. 


sitiens, breeding places of, in 
Lagos, 260, 261, 269; in 
Mauritius, 137. 


sollicitans, breeding in salt 
water in America and India, 
261. 


taeniorhynchus, breeding in 
salt water in America, 254, 
261. 


thalassius, breeding places of, 
in Lagos, 251-253, 260-268. 


(Lutzia) tigripes, in Madagascar 
and Mauritius, 136; breed- 
ing places of, in Lagos, 259- 
269; predaceous larva of, 
281. 

wunivittatus, in Lagos, 269; in 
Madagascar, 137; larva of, 
in Palestine, 389, 394. 


ventrilloni, sp. n., in Madagas- 
ear, 135. 


culiciformis, Anopheles (Stethomyia). 


Culiciomyia nebulosa, breeding places 
of, in Lagos and Gold Coast, 251—270. 


GENERAL 


Culicoides, in North Russia, 197. 
arcuatus, in North Russia, 


39 


198, 

eS fascipennis, in North Russia, 
198. 

9 festivipennis, in North 


Russia, 198. 


pulicaris, in North Russia, 
198. 


cumminsi, Ochlerotatus. 


curta, Phytomia. 

cyanopterun, Rhynchium. 
cyanophylli, Aspidiotus. 

_eycadis, Coccus (see Saissetia oleae). 


Cyathomyia fusca, larva of, at Accra, 
258. 


Cycas revoluta, Coccids on, in Mauritius, 
ia. 


cylindrica, Lachnosterna. 


dactyliperda, Coccotrypes. 


Dacus demmerezi, on Marrows and 
pumpkins in Mauritius, 172. 


»  sygmoides, in Mauritius, 172. 
dahlifolii, Macrosiphum. 
damnosum, Simulium. 
dantini, Dociostaurus. 
Dasyops, 199. 


Dates, pests of, in Mesopotamia, 287— 
303. 


-decens, Culex. 

decisus, Aulacobothrus. 

Deinocerites, in Tropical America, 254, 
WMemmerezi, Dacus. 

denticulata, Lachnosterna. 

depressa, Osca (Pangonia). 


depressum, Lecanium (see Saissetia 
nigra). 

-desertorum, Oryctes. 

destructor, Aspidiotus; Ceroplastes ; 
Tanymecus. 


detritus, Ochlerotatus. 

Dialium guineense (Black Tamarind), 
mosquito larvae breeding in, in 
Lagos, 257. 

Diaspis boisduvali, on Phoenix, 299. 

carueli, parasitised by Aphe- 
linus mytilaspidis in U.S.A., 
297. 

phoenicis, on Phoenix, 299. 

rosae, parasitised by A phelinus 
mytilaspidis in Spain, 297. 


99 


INDEX. 419 
Diatomineura inornata (seeCorizoneura). 
Dicrurus afer, predaceous on Glossina 
in §.E. Africa, 368. 
dictyospermi, Aspidiotus. 
Digera arvensis, food-plant of thrips in 
Sudan, 98. 
digitata, Aclerda (Pseudolecanium). 
dilemma, Lachnosterna. 
dimidiatus, Carpophilus. 
Diplorhynchus mossambicensis, in §.E. 
Africa, 320, 366, 379. 
Dipterocarpus _ turbinatus, 
shoreae on, in India, 277. 
discors, Pangonia. 
Dispholidus, in 8.E. Africa, 379. 
dissimilis, Buplex (Corizoneura). 
distincta, Corizoneura. 
divaricata, Chrysops. 
Dociostaurus, revision of, 397—407.- 
se albicornis, characters 
of, 400, 404. 
x anatolicus, characters of, 
398, 400, 404; dis- 
tribution of, 405. 
¥ crassiusculus, 398; char- 
acters of, 400, 401; 
synonomy of, 405; 
distribution of, 406. 
rs crucigerus, characters of, 
399; synonymy of, 
402; distribution of, 
403, 404. 
crucigerus hispanicus, 
characters of, 399; 
peculiar to Spain, 403. 
crucigerus tartarus, 399, 
403, 404. 
dantini, in Morocco, 397, 
400, 405. 
genei, characters of, 400; 
distribution of, 404. 
gracilis, probably syno- 
nymous with D. genei, 
397, 398. 
kervillei, in Persia, 397. 
kraussi, characters of, 
401; distribution of, 
406, 407. 
kurdus, sp. n., char- 
acters of, 401; dis- 
tribution of, 406. 
maroccanus, distribution 
of, 398, 402; char- 
acters of, 397, 399; 
synonymy of, 401, 
404, 


Calandra 


99 


99 


99 


420 


Dociostaurus plotnikovi, sp. n., charac- 
ters of, 400; synon- 
ymy of, 405 ; distribu- 
tion of, 406. 


< tartarus, 397. 
domestica, Musca. 
domesticus, Ochlerotatus. 


Dominica, new species of Lachnosterna 
from, 191. 


dorsalis, Cerodonta. 


Dracaena fragrans, mosquito larvae 
breeding in, in Lagos, 257 ; 
in relation to distribution 
of Glossina, 319. 


MA gazensis, in S.E. Africa, 320. 
dracaenae, Hretmopodites. 
drapert, Sphaerococcus. 


Dryomyza flaveola, seasonal variation 
or, 214: 


durbanense, Lecanium. 
duttoni, Culex. 


Echidnophaga gallinacea, imported into 
Mauritius, 172. 


egbarum, Ceroplastes. 


Egypt, date pests in, 288, 289, 294; 
mosquitos in, 156, 392, 393, 394, 395; 
Dociostaurus gracilis in, 397. 


ehretiae, Lecanium. 


Eichornia crassipes (Water Hyacinth), 
Anopheles breeding amongst, 73. 


elegans, Oryctes. 

elongata, Pangonia. 
elongatum, Lecanium. 
elytropappt, Inglisia. 
Elytroteinus, nom. n., 276. 


sh subtruncatus, on ginger in 
Fiji and Honolulu, 276. 


Enterolobium dulcis (Rain Tree), mos- 
quito larvae breeding in, in Lagos, 
257. 


Ephestia calidella, in dates in Egypt 
and Mesopotamia, 288. 


Bs cautella, in dates in Meso- 
potamia, 288, 289. 


3 passulella (see E. cautella). 


Epomophorus crypturus, in Glossina 
areas in S.E. Africa, 338. 


equinum, Simulium. 


Erephopsis lata (see Osca). 


GENERAL INDEX. 


Eretmopodiltes conde (see EF, quin- 
quevittatus). 
9 dracaenae breeding 
places of, in Sierra 
Leone, 258. 
‘4 inornatus, in Lagos, 269. 
> quinquevittatus, breeding 
places of, in Lagos, 
264, 265, 270; in 
Madagascar, 138. 
Eriodendron orientale (Silk Cotton Tree),. 
mosquitos breeding in, in Lagos, 256. 


Eritrea, Lonchaea in, 201, 203. 

Erythroxylon 
Africa, 322. 

eucleae, Ceroplastes. 


Eugenia, in relation to breeding places. 
of Glossina morsitans in 8.E. Africa, 
364. 


exaptus, Systates. 


emarginatum, in §.E. 


excisa, Lonchaea (Carpolonchaea). 
expansum, Lecanium. 
extricator, Culex. 


fasciata, Stegomyia. 
fascipennis, Culicoides. 
fatigans, Culex. 

feae, Gagrella. 


| ferrugineus, Rhynchophorus. 


fervida, Lachnosterna. 

festivipennis, Culicoides. 

Ficalbia neireti (see Uranotaenia). 
ficiperda, Lonchaea (Carpolonchaea). 


| Ficus (Banyan), mosquito larvae’ 
breeding in, in Lagos, 257, 
258. 


»  subcalearata, in 8.E. Africa, 323. 
jficus, Aspidiotus. 
Figs, bionomics of Ocnerogyia amanda: 
on, in Mesopotamia, 181-186. 
Fiji, new weevils from, 276. 
filamentosum, Lecanium. 
Filaria bancrofti, carried by mosquitos. 
in Lagos, 255. 
jfilicum, Lecanium hemisphaericum (see 
Saissetia). 
filiformis, Ischnaspis. 
Filippia, 2, 23. 
as africana (see F. carissae). 


Ae carissae, sp. 0., on Carissa 
grandiflora in Natal, 24. 


* chilianthi, sp. n., in Natal, 23.. 


GENERAL INDEX. 


Fiorinia pellucida, on Phoenix, 299. 


flava, Myzomyia (see Anopheles wm- 
maculatus). 


flaveola, Dryomyza. 
jflaveolum, Lecanium (see L. hesperidum). 


flavus, Taeniorhynchus (Culex, Coquil- 
lettidia). 


floccifera, Pulvinaria (Coccus). 

jloccosa, Pulvinaria (see P. floccifera). 

fluviatilis, Aédes (Culex). 

Forda natalensis, sp. n., in Natal, 71. 
h rotunda, 72. 


Formaldehyde, as a poison for house- 
flies, 47-63 ; impurities in, 47. 


Formic Acid, disadvantages of, in 
formaldehyde, 50. 


formosa, Corizoneura. 


fowleri, Culex (see Ochlerotatus nigeri- 
ensis). 


fragilis, Anopheles (see A. algeriensis). 


France, Aphelinus mytilaspidis para- 
sitic on Parlatoria blanchardi in, 297. 


fraseri, Uranotaenia bilineata. 
fryeri, Aédes (Ochlerotatus). 
Fryi, Stenommatus. 


Fuirena, in relation to distribution of. 


Glossina, 326, 345. 
fuliginosus, Anopheles (Nyssorhynchus). 
fulvifascia, Osca (Pangonia). 
funestus, Anopheles. 
fusca, Cyathomyia. 
fuscinervis, Buplex. 
fuscopennatus, Taeniorhynchus. 


fuscus, Pectinopalpus (see Ouliciomyia 
nebulosa) ; Thriambeutes. 


Gagrella feae, an imported pest in 
Mauritius, 174. 


Galago crassicaudatus, occurrence of, in 
Glossina areas in §.E. Africa, 337. 


gallinacea, Echidnophaga. 
Game, in relation to Glossina in Africa, 


115, 117, 327, 328, 333, 336-339, 342,. 


353, 376. 
gasteralpha, Pulvinaria. 


Gelechiid, unidentified species of, on 
date palms in Mesopotamia 290, 303. 


genet, Dociostaurus (Stauronotus). 
geniculatus, Ochlerotatus. 
geranui, Inglisia. 


421 


giganteus, Culex. 
glandium, Calandra. 


Glossina austeni, bionomies of, in S.E. 
Africa, 329, 330, 348, 349, 
350, 351, 366. 


a brevipalpis, bionomics of, in 
Nyasaland, 102; bionomics. 
of, in 8.E. Africa, 315-384. 


by longipalpis, 339. 


o morsitans, bionomics of, in 
S.E. Africa, 315-388; in 
Nyasaland, 101 ; bionomics 
of, in 8. Rhodesia, 113-132. 


pallidipes, bionomics of, in 
S.E. Africa, 316—339. 


»  palpalis, in Uganda, 45. 
glossinae, Mutilla. 
Glugea, parasite of Simulium, 215. 


Gold Coast, Lonchaea impressifrons in,. 
202; mosquito larvae from, 105- 
1 


gracilis, Dociostaurus. 

grahami, Culex. 

graminis, Chionaspis. 

grandidieri, Culex (see Taeniorhynchus 
flavus). 

guiarti, Culex. 

Guttera edouardi, possibly destroying 


pupae of Glossina in §.E. Africa, 328, 
369. 


) 


Habrobracon kitcheneri, parasite of 
Ephestia cautella, in Egypt, 289. 


Haematopota, in North Russia, 196, 


197. 

es crassicornis, in North 
Russia, 198. 

ie pluvialis, in North 


Russia, 198. 
haemorrhoidalis, Heliothrips. 
hastata, Oorizoneura. 
hauensteini, Dociostaurus (Stauronotus) 
(see D. crassiusculus). 
Hawaii, new weevils from 276, 277. 
hebraeus, Polistes. 
hederae, Aspidiotus. 
Hedychium coronarium (Ginger), weevil 
on, in Hawaii, 276. 
Heliothrips haemorrhoidalis, on date 
palm in Trinidad, 295. 


99 indicus (Cotton Thrips), 
bionomics and control 
of, in Sudan, 95-100. 


422 


Heliotropium europaeum, food-plant of 
thrips in Sudan, 98. 


si supinum, food-plant of 
thrips in Sudan, 98. 


Hemilecanium, 2, 14. 


a theobromae, on Huphor- 
bia and oleander in 
Cape Colony, 14. 


hemisphaerica, Saissetia. 
Heptaphlebomyia argenteopunctata (see 


Culex). 
59 kingi (see Culex 
argenteopunctatus). 
4 montforti (see Culex 
uniwittatus). 


hesperidum, Lecanium (Cocecus, Calym- 
natus). 


Hexamethylene Tetramine, influence 
of, in formaldehyde, 48. 


hibernaculorum, Lecanium (see Sais- 
setia hemisphaerica). 


hirtipes, Simulium. 

hispanicus, Dociostaurus crucigerus. 
histrio, Oedipoda. 

Hodgesia sanguinis, in Lagos, 269, 270. 


Holotrichia, considered a synonym of 
Lachnosterna, 189. 


Hopperdozer, new type of, 179. 

hornigu, Spermatophora. 

hortensis, Culex. 

House-flies, formaldehyde as a poison 
for, 47-63. 

Hyalopterus arundinis, on Vitis, 66. 

hydatis, Coccus (Calypticus) (see Cero- 
plastes rusci). 

Hylochaerus, possible relation of, to 
Glossina in 8.E. Africa, 337. 


Hyphaene  ventricosa, puparia of 
Glossina morsitans found near, in 
S.E. Africa, 364. 

Hypocolius ampelinus, injuring dates 
in Mesopotamia, 301. 

hyreanus, Anopheles. 


Icerya aegyptiaca, on date palm in 
Zanzibar, 299. 


ay purchasi, on date palm, 299. 
Idiosaissetia, gen. n., 2, 40. 
” peringueyt, sp. ., on 
grass in 8. Africa, 40. 
ammaculatus, Anopheles (Myzomyia). 
ampressifrons, Lonchaea. 


GENERAL INDEX. 


inafectatus, Tanymecus. 

inconspicuosus, Micraedes. 

indefinitus, Anopheles (Myzomyia) (see 
A. vagus). 


India, pests of date palms in, 293, 294, 
295; Lonchaea spp. in, 206, 208, 
209; Anopheles breeding in salt 
water in, 261; new weevils trom, 
276, 277. 


indicus, Heliothrips. 
Inglisia, 2, 26. 


= elytropappi, sp. n., on Elytro- 
pappus rhinocerotis in Cape 
Colony, 36. 


cs geranii, sp. D., on geranium 
in Cape Colony, 37. 


9 zizyphi, sp. n., on Zizyphus in 
Transvaal, 37. 
ingrami, Phlebotomus. 


insulae-florum, Anopheles (Stethomyia) 
aitkent. 


inornata, Bouvierella; Cort.on ura 


(Diatomineura). 
inornatus, Hretmopodites. 
insatiabilis, Culex. 
insignis, Culex. 
interpunctella, Plodia. 
invidiosus, Culex (see C. decens). 


Ipomoea cordofana, food-plant of thrips 
in Sudan, 98. 


irritans, Ochlerotatus. 


Isaniris ater, sp. n., on Brachystegia in 
Nyasaland and Rhodesia, 273. 


Ischnaspis filiformis, on Phoenix, 299. 

Isoglossa mossambicensis, in 8.E. Africa, 
o22. 

italicus, Calliptamus. 

Italy, Ochlerotatus ornatus in, 259 . 
Aphelinus mytilaspidis parasitising 
Parlatoria blanchardi in, 297. 

jacksom, Pulvinaria. 

Jamaica, Lachnosterna in, 190, 191. 

jamaicensis, Lachnosterna. 

Japan, Nippolachnus piri in, 157. 


Java, Anopheles in, 77; Lonchaea in, 
207, 209. 


kellyi, Saissetia. 


kervillei, Dociostaurus (see D. crasstus- 
culus). 


GENERAL INDEX. 


Khaya nyasica (East African Mahog- 
any), relation of Glossina to, in S.E. 
Africa, 319, 320, 346, 348, 349, 376. 


Kigelia pinnata, relation of Glossina to, 
in S.E. Africa, 348. 


Killing-bottle, description of new type 
of, 283-285. 


kingi, Heptaphlebomyia 
argenteopunctatus). 


kitcheneri, Habrobracon. 
kochi, Anopheles (Cellia). 
kraussi, Dociostaurus. 
kurdus, Dociostaurus. 


(see Culex 


Lachnosterna, synonyms of, 189. 


a acinosa, Sp. 0., in 
Trinidad, 190, 192. 

a anodentata, in Mexico, 
192. 

eS antiguae, sp. n., in 


Antigua and Domin- 
ica, 190, 191. 


- (Phytalus) cometes, 193. 

s cylindrica, 190. 

= denticulata, in Mar- 
tinique, 192. 

a dilemma, sp. n., in Trini- 
dad, 190, 192, 193. 

a fervida, in Jamaica, 190. 

is jamaicensis, Sp. N., in 
Jamaica, 190, 191. 

e longipilosa, in Mexico, 
192. 

be misteca, in Mexico, 192. 

Wy montserratensis, Sp. D., 
in Montserrat, 190, 
191. 

ov (Phytalus) omiltemus, in 
Mexico, 189. 

is (Phytalus) platyrrhinus, 
in Mexico, 189. 

. setifera, in Central 
America, 193. 

ey (Phytalus) smithi, bion- 


omics and control of, 
in Mauritius, 159-169, 


P7E, bie ae’ * Weat 
Indies, 189. 

a trinitatis, in ‘Trinidad, 
190, 193. ~ 


Lachnus pyri, on pears in Persia, 156. 


Laemophloeus sp., infesting stored dates 
in Mesopotamia, 303. 


423 


laevis, Calypticus (see Lecanium hespert- 
dum); Lonchaea (Carpolonchaea). 


Lagerstroemia regina, mosquito larvae 
breeding in, in Lagos, 257. 


Lagos, breeding places of mosquitos 
in, 247-270. 


lamberti, Stegomyia (see S. albopicta). 
lambiana, Lonchaea. 
Landecrab (see Cardisoma). 


Landolphia, relation of Glossina to, 
in §.E. Africa, 317, 320, 322, 333. 


lata, Osca (Hrephopsis). 
lataniae, Aspidiotus. 
lateralis, Corizoneura. 
laticinctus, Culex. 
latipes, Simulium. 
laurenti, Culex. 
laurt, Chermes 
peridum). 

lautissima, Pangonia. 

Lecaniodiscus fraxinifolia, in relation 


to distribution of Glossina in S.E. 
Africa, 349. 


(see Lecanium hes- 


Lecanium africanum, on citrus in 

Natal, 4. 

25 alienum (see L. hesperidum). 

# angustatum (see LL. hes- 
peridum). 

at artemisiae (see Ceroplastes 
TUSC1). 

"p assimile var. amaryllidis 

(see L. hesperidum). 

- beaumontiae (see Saissetia 
hemisphaerica). 

és begoniae (see Saissetia nigra). 

5 cassinae (see Saissetia oleae). 

x clypeatum (see Saissetia 
hemisphaerica). 

+ coffeae (see Saissetia hemi- 
sphaerica). 

és depressum (see Saissetia 
nigra). 

es durbanense, sp. n., in Natal. 

ye ehretiae, sp. n., on Ehretia 
hottentottica in Trans- 
vaal, 4. 

" elongatum, on Acacia melan- 
oxylon in Transvaal, 5. 

59 expansum, on Phoenix, 299. 

= filamentosum, on fig in Cape 
Colony, 6. 


424 
Lecanium flaveolum (see DL. hespert- 
dum). 
a hemisphaericum (see Sais- 
setia). 
55 (Coccus) hesperidum, on 


date palms in California, 
299; on citrus in Cape 
Colony, 3; on limes in 
Mauritius, 173, 174. 
9 maculatum (see L. hespert- 
dum). 
mangiferae, food-plants of, 
in Mauritius, 173, 174. 
minimum (see L. hesperi- 
dum). 
nanum (see IL. hesperidum). 
2s nigrum (see Saissetia nigra). 
he oleae (see Saissetia). 


AS persimile (see Saissetia per- 
similis). 

» platycerii (see L. hesperi- 
dum). 


a proteae, sp. n., on Protea in 
Transvaal, 7. 


4 pseudelongatum, on Acacia 
caffra in Transvaal, 6. 

x pumilum, sp. 1., in Cape 
Colony, 5 

re signiferum (see L. hesperi- 
dum). 

By subpatelliforme (see Sais- 
setia). 

se sylvestru (see Saissetia per- 
similis). 

Bs terminaliae (see L.  hes- 
peridum). 

™ tessellatum, on Phoenix, 299. 

x ventrale (see L. hesperidum). 

b> wistariae, Sp. D., on Wis- 


taria in Cape Colony, 8 
Lemna, Anopheles breeding amongst, 
74. 


lepida, Pulvinaria. 


Lepidosaphes ulmi, parasitised by 
Aphelinus mytilaspidis in U.S.A., 
297. 


Leucas nubica, food-plant of thrips in 
Sudan, 98. 

Jleucosphyrus, 
mya). 

Leva apicalis, synonymy of, 398. 
» mundus, synonymy of, 398. 


Lichtensia asparagi, sp. n., on Aspara- 
gus capensis in Cape Colony, 23. 


Anopheles (Neomyzo- 


GENERAL INDEX. 


lineatithoraz, Corizoneura. 
lineatopennis, Banksinella. 


Lispa, bionomics of, in Nyasaland, 
279-281. 

lividocostalis, Culex. 

lloydi, Villa. 

Lonchaea, key to Ethiopian species of, 
199, 200; key to Indo- 
Australian species of, 
204; key to Oriental 
females of, 210. 


» (Lamprolonchaea) aurea, 
wide distribution of, 200, 
202, 204, 206. 


7” (Carpolonchaea) atratula, in 
Malaya and Philippines, 
205,207. 


‘3 biroi, probably a synonym 
of L. atratula, 205, 207. 


9 (Carpolonchaza)  bisulcata, 
sp. n., in India, 206, 208, 
210. 


mA brouniana (see L. aurea). 


f (Carpolonchaea) citricola, on 
oranges in Philippines 
and Australia, 205, 207. 


a continentalis, sp. ., in 
Natal, 200, 202. 


> (Carpolonchaea) excisa, wide 
distribution of, 201, 206, 
209, 210. 


> (Carpolonchaea) ficiperda, in 
Philippines, 205, 207, 210. 


as impressifrons, sp. in 
Gold Coast, 201, 209. 


” Soe gee ig laevis, sp. 
.,in Zanzibar and Eri- 
my 201, 203. 


Pe lambiana, in Seychelles, 200, 
202. 

> (Carpolonchaea) lucens, in 
Java, Sumatra and Philip- 
pines, 206, 209, 210. 

wis metatarsata (see D. aurea). 


” minuta, in Java and Philip- 
pines, 205, 207. 


¥ mochii, from Eritrea, 200, 
201. 

“ obscuripennis, probably a 
synonym of L. atratula, 
207. 


* (Carpolonchaea) ophyroides, 
sp. n., in Tropical Africa 
201, 203. 


oo, ___——< = 


—— —— ——_ —EE— 3 a a Se “2 — 7 


6 tie 
- _ = 


GENERAL INDEX. 425 


Lonchaea (Carpolonchaea) pallicarpa, 


sp. n., in Philippines, 205, 
207, 210. 
a (Dasyops) phaeolepis, sp. 0., 
in Natal, 200, 201. 
(Carpolonchaea) plumosis- 
sima, wide distribution 
of, 199, 201, 203, 204, 
206, 209, 210. 
(Carpolonchaea) setifera, in 
New Guinea and Philip- 
pines, 206, 209, 210. 
= splendida (see L. awrea). 


Lonchaeidae, classification of new 
African and Oriental species of, 199- 
210 


longiareolata, Theobaldia. 
longicorda, Ceroplastes. 


longipalpis, Glossina; Ochlerotatus ; 
Phlebotomus. 


99 


99 


longipilosa, Lachnosterna. 
longirostris, Corizoneura. 


Loranthus, bird-lime made from, used 
in catching tsetses, 361. 


dlounsburyi, Oryptinglisia. 
ducens, Lonchaea (Carpolonchaea). 


ludlowi, Anopheles (Myzomyia). 
luridus, Tabanus. 


dutescens, Ochlerotatus. 
luteocephala, Stegomyia. 


LIutzia, 136. 


Lygodium subalatum, in S.E. Africa, 


322. 


-maculatum, Lecanium (see L. hes- 


peridum) ; Simulium. 
maculicornis, Tabanus. 


-maculipennis, Anopheles. 


Macrosiphum cissi, sp. n., on Cissus in 
Transvaal, 65. 


ys dahliafolii, on Carduus 
in Uganda, 72. 

; sonchi, on Carduus in 
Uganda, 72. 

a viticola, on Vitis, 66. 


“Madagascar, mos uitos of, 113-138. 


‘madagascariensis, Pseudoheptaphlebo- 


myia (see Culex quasigelidus). 
Malata, food-plant of thrips in Sudan, 
98. 


“Malaya, Anopheles in, 77-92; Lonchaea 
spp. in 204, 207, 209. 


Malapterurus electricus (Electric Cat- 


fish) feeding on mosquito larvae in | 


Lagos, 266. 


Mangifera indica (Mango), mosquito 
larvae breeding in, in Lagos, 257 ; 
Coccids on, in Mauritius, 174. 

mangiferae, Lecanium. 


Mansonia titillans, breeding among 
water lettuce, 73, 75. 


Mansonioides africanus, breeding places 
of, in West Africa, 
14, 258; 266,268, 
269, 270. 


5 uniformis, in Lagos, 269 ; 
in Madagascar, 135. 


mariae, Ochlerotatus. 


Markhamia, puparia of Glossina au tent 
found under logs of, 366. 
ef lanata, in relation to dis- 
tribution of Glossina in 

S.E. Africa, 348, 382. 


marlatti, Phoenicococcus. 
maroccanus, Dociostaurus. 
marshali, Anopheles ; Ochlerotatus. 


Martinique, Lachnosterna denticulata 
in’ 192. 


mashonaensis, Uranotaenia. 
mauritianus, Anopheles ; Aspidiotus. 


Mauritius, mosquitos of, 113-138; 
bionomics and control of Lachnos- 
terna (Phytalus) smithi in, 159-169 : 
plant pest legislation in, 175-177; 
insects accidentally introduced into, 
171-177 ; new weevil from, 276. 


melanopyga, Phara (Pangonia). 
Mellera lobulata, in 8.E. Africa, 322. 
Membranaria, gen. n., 2, 41. 


“ pretoriae, Sp. D., on grass 
in Transvaal, 41. 


meridionalis, Pericoma. 

Mermis, parasite of Simuliwm, 215. 
mesambrinoides, Pangonia. 
mesembryanthemi, Pulvinaria. 


Mesopotamia, bionomics of Ocnerogyia 
amanda on figs in, 181-186; date 
pests in, 287-303; Phlebotomus 
sergenti var. in, 310, 


mespili, Myzus. 

mespiliella, Myzus. 
Mespilus, new Aphid on, 155. 
metallica, Stegomyia. 
metallicus, Taeniorhynchus. 


Metarrhizium anisopliae, introduction 
of, into Mauritius from Porto Rico, 
166. 


426 


metatarsata, Lonchaea (see L. aurea). 

Methylamine, influence of, in formalde- 
hyde, 48. 

Mexico, Lachnosterna spp. in, 189, 192. 


Micraédes <inconspicuosus, breeding 
places of, in Lagos, 251-270. 


Milletia sanagana, mosquito larvae 
breeding in, in Lagos, 257. 


»  stuhlmanni, in S.E. Africa, 


321. 
mimeticus, Culex. 
Mimomyia mimomyiaformis, in Lagos, 
269. 
splendens, breeding among 


water-lettuce in West 
Africa, 74. 


mimomytaformis, Mimomyia. 
mimosae, Ceroplastes. 


minimum, Lecanium (see L. hesperi- 
dum). 


minimus, Anopheles (Myzomyia). 

minuta, Lonchaea. 

minutus, Ochlerotatus ; Phlebotomus. 

mirandum, Lecanium oleae (see Sais- 
setia). 

misteca, Lachnosterna. 

mobilis, Ceronema. 

mochii, Lonchaea. 

modestus, Culex. 


moluccensis, Anopheles (Nyssorhynchus) 
annulipes. 


montforti, Heptaphlebomyia (see Culex 
wunivittatus). 
monticolor, Simulium. 


Montserrat, new Jachnosterna from, 
191. 


montserratensis, Lachnosterna. 
Morocco, Dociostaurus in, 401, 405. 


39 


morsitans, Glossina; Simulium ; Theo- 
baldia. 


Mosquitos, breeding places of, in West 
Africa, 105—112, 247-270; in Mada- 
gascar, 133-138; Lispa predaceous 
on, in Nyasaland, 279-281 ;. biono- 
mics of, in Palestine 387-395. 


multicolor, Anopheles. 
mundus, Leva (Stenobothrus). 


Mungos persicus, possibly destructive 
to dates in, Mesopotamia, 300. 


musae, Stenommatus. 


GENERAL INDEX. 


Musca domestica, 368; formaldehyde 
aS a poison for, 47. 


Mutilla benefactrix, parasite of Glossina 
morsitans, in Nyasaland, 104. 


»  glossinae, parasite of Glossina 
morsitans in Nyasaland, 101. 


Myelois phoenicis, in dates in Algeria, 
288. 


Myosorex, in Glossina areas in S.E.~ 
Africa, 338. 


myricae, Ceroplastes (Coccus). 

mytilaspidis, Alphelinus. 

Myzomyia flava (see Anopheles wm- 
maculatus). 

Myzus mespili, on medlar, 155. 


i mespiliella, sp. n., on medlar in. 
Persia, 154. 


Nagana, recent outbreaks of, in 8.E- 
Africa, 372. 


nanum, Lecanium (see L. hesperidum).. 

Natal, Lonchaea spp. in, 201, 202. 

natalensis, Forda. 

neavei, Corizoneura. 

nebulosa, Culiciomyia. 

neireti, Culex ; Uranotaenia (Iicalbia).. 

Newbouldia laevis, mosquitos breeding 
in, in Lagos, 257. 

New Guinea, Lonchaea in, 207, 209; 
Anopheles in, 81-83. 
Newsteadina arboricollis 

podomyia). 

Nigeria, Lonchaea ophyroides in, 204 
mosquito breeding places in, 247— 
270. 

nigeriensis, Ochlerotatus (Aédes). 

nigerrimum, Tapinoma erraticum. 


(see Ortho- 


nigra, Saissetia. 

nigrella, Sarssetia. 

nigricephalus, Ochlerotatus. 

nigricornis, Tabanus. 

migrifacies, Tabanus.« 

nigripes, Anopheles ; Ohrysops. 

nigrocostalis, Culex. 

nigro-emarginatus, Lasius emarginatus. 

nigrum, Lecaniwm (see Saissetia nigra). 

nili, Anopheles. 

nipae, Pseudococcus. 

Nippolachnus piri, on Pyrus sinensis in 
Japan, 157. 


——_— 


GENERAL INDEX. 


Numida mitrata, relation of, to Glos- 
sina in §.E. Africa, 328, 369. 


nurset, Anopheles (see A. swperpictus). 


Nyasaland, bionomics of Glossina in, 
101-104; bionomics of predaceous 
Dipteron in, 279-281; new weevil 
from, 273. 


obreptans, Simulium. 
obscuratus, Anopheles. 
obscuripennis, Lonchaea. 


Ocimum basilicum, food-plant of thrips 
in Sudan, 98. 


Ochlerotatus alpinus, in Norih Russia, 

198. 

= apicoannulatus, breeding 
places of, in Lagos, 256, 
258, 270. 

os argenteopunctatus, in Lagos, 
268, 270. 

zs caliginosus, breeding places 
of, in Lagos, 253, 268, 
270. 

a caspius, in Egypt and 
Palestine, 393. 

= cumminst, in Lagos, 268, 
270. 

of detritus, in Palestine, 393. 

a domesticus, in houses in 
Lagos, 267, 268, 269, 
270. 

ry fryeri, 134. 

_ geniculatus, breeding places 
of, in England, 259. 


Pe irritans, breeding places 
of, in Lagos, 251-270. 

.r longipalpis, breeding places 
of, in Lagos, 256, 269. 

lutescens, in North Russia, 
198. 

E mariae, in Palestine, 393. 

+ marshalli, breeding places 
of, in Lagos, 256, 269. 

a minutus, breeding places 
of, in Lagos, 258. 

2 nigeriensis, 1n Mauritius, 
134. 

“a nigricephalus, breeding 
places of, in Lagos, 251— 
270. 

a ochraceus, in Lagos, 269, 
270 

f ornatus, breeding i in horse- 
chestnut in Italy, 259. 

Pe punctothoracis, 137; in 


Lagos, 253, 268, 270. 
(741) 


427 


ochraceous, Ochlerotatus. 
oculata, Saissetia, 


Ocnerogyia amanda, bionomics on om 
figs in Mesopotamia, 181, 185, 186. 


Oedipoda brevicollis (see Dociostauwrus 
crucigerus). 


a histrio, probably a form of 
Calliptamus italicus, 398. 


Oenanthe, Simulium breeding amongst, 
224. 

oldi, Pangonia. 

oleae, Saissetia (Coccus). 

oleracea, Tipula. 


Oligonychus (Paratetranychus) simplex, 
control of, on date palms in Meso- 
potamia, 299, 300, 303. 


omiltemus, Lachnosterna (Phytalus). 
ophyroides, Lonchaea (Carpolonchaea). 
Orange (see Citrus aurantium). 
orientalis, Vespa. 


Ormosia angolensis, relation of, to 
Glossina morsitans in S.E. Africa 
365. 


ornata, Uranotaenia. 
ornatum, Simulium. 
ornatus, Ochlerotatus. 


Orthocladius, parasite of Simulium, 
215. 


Orthopodomyia arboricollis, in Mauritius, 
135. 


Oryctes desertorum, distribution of, 293, 
294. 


, elegans, on date palms in 
Mesopotamia, 292. 


»  Thinoceros, on date palms in 
India ete., 293, 294. 


Osca barbata, 139. 
» (Pangonia) depressa, 139. 
»  fulvifascia, 139. 
» lata, synonymy of, 139. 
ostreaeformis, Aspidiotus. 


Otomys, in Glossina areas in 8.E. Africa, 
338. 


Oxyanthus gerrardi, in 
distribution of Glossina, in 
Africa, 349. 


relation to 
S.E. 


pacificum, Lecanium (Calymnatus) (see 
L. hesperidum). 


Padraona palmarum, on date palms in 
Caleutta, 288. 


Cc 


-428 


Palestine, bionomics of mosquitos in, 
387-395. 


- palestinensis, Anopheles (see A. super- 
pictus). 

pallicarpa, Lonchaea (Carpolonchaea). 

pallidipes, Glossina. 

pallidipennis, Corizoneura. 


pallidus, Anopheles (Myzorhynchus) 
barbirostris ; Ceroplastes. 


palmae, Coccus (see Saissetia oleae) ; 
Parlatoria proteus. 


palmarum, Padraona. 
palpalis, Glossina. 
paludosa, Tipula. 


Panama, Anopheles in, 73-75; Aédes 
fluviatilis breeding in salt water in, 
261. 


Pandanus pacificus (Broad-leaved Screw 
Pine), mosquito larvae in, 
in Lagos, 257. 


a veitchi, mosquito larvae 
in, in Lagos, 256, 257. 
Pangoma appendiculata (see Cori- 
zoneura aethiopica). 
=F barbata (see Osca). 
a beckert, in East Africa, 148, 
149. 
5 brunnipennis (see Buplex). 
a carpenteri, sp. n., in Tan- 
ganyika Territory, 149, 151. 
- discors, sp. n., in Angola, 
148, 149. 
as elongata, 149. 
a fulvifascia (see Osca). 
35 lautissima, sp. n., in Tan- 
ganyika Territory, 148. 
a: mesambrinoides, incorrectly 


placed in this genus, 149. 
- oldi, 145, 146. 


as rufa (see Corizoneura tapro- 
banes). 


a suavis (see Buplex). 
os subfascia (see Buplez). 


ws zonata, Oorizoneura inornata 
recorded as, 145, 146. 


papatasii, Phlebotomus. ' 
pupuae, Anopheles (Stethomyia) aitkeni. 
Parafairmairea, 2, 39. 

ag patellaeformis, sp. Di, 


on Acacia karroo in 
Cape Colony, 39. » 


GENERAL INDEX. 


parallela, Tiphia. 


Paratetranychus simplex (see Oligony- 
chus). 


Parkia biglobosa, mosquitos breeding 
in, in Gold Coast, 259. 


Parlatoria blanchardi, on date palms, 
etc., 295, 296, 297, 299; 
control of, 297; para- 
sites of, 296, 297. 


- pergandei, on Phoenix, 299. 


proteus var. palmae, on 
Phoenix, 299. 
Be victrix (see P. blanchardt). 


= zizyphi, on date palms, 298, 
299. 


Passer domesticus, destructive to dates 
in Mesopotamia, 301. 


»  hispaniolensis, destructive to 
dates in Sudan, 301. 


passulella, E’phestia. 
patellaeformis, Parafawmairea. 


patelliformis, Coccus (see Lecanium 
hesperidum). 


pectinopalpus fuscus (see Culiciomyia 
nebulosa). 

peditaeniatus, 
chus). 

pellucida, Fiorinia. 

pembaensis, Aédes (Skusea). 

penetrabilis, Corizoneura. 

penetrans, Sarcopsylla. 

peregrina, Schistocerca. 

pergandei, Parlatoria. 

Peritymbia vitisana, on Vitis, 66. 

Pericoma meridionalis, in Lagos, 262. 


Anopheles (Myzorhyn- 


peringueyt, Idiosaissetia. 
permutans, Chionaspis. 
perniciosus, Phlebotomus. 


Persea gratissima (Avocado Pear), 
Coccids on, in 8. Africa, 11, 
ie 


perseae, Saissetia. 


Persia, new Aphids from, 153-157; 
Phlebotomus sergenti in, 307 ; Oryctes 
desertorwm in, 294; Dociostaurus 
maroccanus in, 402. 

persimilis, Saissetia. 

Petrodomus, believed to destroy Glossina 
pupae in S.E, Africa, 338, 369. 


phaeolepis, Lonchaea (Dasyops). 


GENERAL INDEX. 


phaiae, Pulvinaria (see P. floccifera). 
Phara, Cadicera a synonym of, 140. 
o (Pangonia) chrysostigma, 140. 
a (Pangonia) crassipalpis, 140. 
os (Pangonia) melanopyga, 140. 
A (Cadicera) rubramarginata, 140. 
pharoensis. Anopheles. 


Pheidole cockerelli, associated with 
Coeccids in Arizona, 296. 


Philippines, Lonchaea in, 207, 209. 


Phlebotomus, not found in North 
Russia, 197; classifi- 
cation of, 305-311. 


antennatus, male of, 305. 


- caucasicus (see PP. ser- 
genti). 

ea ingrami, in Gold Coast, 
306. 

% longipalpis, distinct from 
P. walkeri, 311. 

“- minutus, distribution of, 
306. 

<3 minutus africanus, dis- 


tribution of, 306. 


papatasti, distribution of, 
~ 306, 307, 308. 


ae perniciosus, distribution 
of, 307. 
ot sergenti, in Persia, 307, 


308, 309, 310; variety 
of, in Mesopotamia, 


309. 

i signatipennis, Sp. 0., in 
Gold Coast, 310. 

a similimus, distribution 
Vhs) 00; 300... 

i walkeri, distinct from 


| P. longipalpis, 311. 
phoenicis, Diaspis; Myelois; Rhyn- 
chophorus. 


Phoenicococcus, on date palms, 
297. 


3g marlatti, 297; bion- 
omies and control 
of, 298, 299. 


Phoenix dactylifera (see Date Palm). 


Phorodon violae, sp. n., on pansy in 
Natal, 67, 68. 
Phyllanthus niruri, food-plant of thrips 
in Sudan, 98. 
Phyllastrephus capensis, relation of, to 
Glossina in S.E. Africa, 369. 
(741) 


296, 


499 


Phyllophaga (see Lachnost rn1). 
Phylloxera, on Vitis, 66. 


Phytalus, considered a synonym of 
Lachnosterna, 198, q.v. 


Phytomia curta, attacked by mosquito 
larvae in Nyasaland, 281. 


pinico'a, Lecanium minimum 
L. hesperidum). 


pinifolii, Chionaspis. 
pipiens, Culex. 


Piptadenia buchanani, in relation to 
distribution of Glossina in 8.E. Africa 
319, 322. 


piri, Nippolachnus. 

piriformis, Protopulvinaria. 
piscicidium, Simulium. 
Pistia stratiotes (Water-lettuce), re- 


lation of, to mosquito larvae, 73, 
253. 


Plant Pest Legislation, in Mauritiu-, 
165, 175. 


platycerti, Lecanium (see L. hesperidum). 
platyrrhinus, Lachnosterna (Phytalus). 


Pleuroridgea zanguebarica, in 8.E. 
Africa, 322. ° 


Plodia interpunctella, 
Arizona, 288. 


plotnikovi, ‘Dociostaurus. 
plumbeus, Anopheles. 


plumosissima, Lonchaea 
chaea). 


pluvialis, Haematopota. 

Poinciana regia (Flamboyant), mos- 
quitos breeding in, in Lagos, 257. 

Polistes hebraeus, on diseased dates in 
Mesopotamia, 300. 

pollinctor, Stegomyia (see Ochlerotatus 
longipalpis). 


(see 


in dates in 


(Carpolon- 


Polygala triflora, food-plant of thrips 
in Sudan, 98. 

pomorum, Mytilaspis (see Lepidosaphes 
ulm). 


Potamogeton, Simulium _ breeding 
amongst, 224, 237. 

pretoriae, Membranaria. 

Procavia, in Glossina areas in S.E. 
Africa, 338. 


Proceras sacchariphaga, in Mauritius, 
i 
proteae, Lecanium. 
c2 


430 


proteus var. palmae, Parlatoria (see 
P. blanchardi). 


Protopulvinaria, 2, 16. 


os piriformis, on avocado 
pear in Natal, 17. 


pruina, Culex. 

pseudelongatum, Lecanium. 
Pseudococcus nipae, on date palm, 299- 
Pseudolecanium digitatum (see Aclerda). 


Pseudoheptaphlebomyia madagascarien- 
sis (see Culex quasigelidus). 


pseudopictus, Anopheles (see A. hyr- 
canus). 

psidu, Pulvinaria. 

Psidium quayava (Guava), mosquitos 
breeding in, in Lagos, 257. 

Pteleopsis myrtifolia, in S.E. Africa, 
ak: 


Pterocarpus spp., in S.E. Africa, 320, 
322, 323, 349, 351, 352, 383. 


Pteroporus, (see Elytroteinus). 
pulicaris, Culicoides. 
Pulvinaria, 2, 17. 


_ antigoni, food-plants of, 
in Mauritius, 173, 174. 

oa aristolochiae, on Hrythrina 
caffra in Natal, 18. 

Sh brassicae (see P. floccifera). 

i floceifera, on Solanum in 
Natal, 19. 

PA floccosa (see P. floccifera). 

Ss gasteralpha, in Mauritius, 
el. 

ms jacksoni, on Trichilia sp. 
in Natal, 19. 

a lepida, on grass in Trans- 
vaal, 20. 

v5 mesembryanthemi, on Me- 


sembryanthemum in Cape 
Colony, 20. 


» phaiae (see P. floccifera). 


9 psidii, on guava in Natal, 
21. 


pumilum, Lecaniwm. 

punctocostalis Banksinella. 

punctothoracis, Ochlerotatus (Aédes). 

punctulatus, Anopheles (Nyssorhynchus) 

Punica granatum, Aphids on, in Italy 
and Egypt, 156. 

» sylvestris, Aphis punicae on, in 

Italy, 156. 


GENERAL INDEX. 


punicae, Aphis. 
punicellae, Aphis. 
purchasi, Icerya. 


Pycnonotus leucotis, destructive to 
dates in Mesopotamia, 301. 


pyrt, Lachnus. 
Pyrus sinensis, Aphid on, in Japan, 157 


quadrilineatus, Ceroplastes. 
quasigelidus, Culex. 


Quercus dilatata, Calandra glandiwm in 
acorns of, in India, 277. 


aA imeana, new weevil in acorns 
of, in India, 277. 


quinquevittatus, Hretmopodites. 


radiatus, Oalypticus (see Ceroplastes 
TUSsCt). 
Ranunculus, Simulium breeding 


amongst, 213, 241. 
rapax, Aspidiotus. 
Rauwwolfia inebriens, in 8.E. Africa, 321. 
relicta, Chrysops. 
reptans, Simulium. 
reticulatus, Culex. 
Réunion, mosquitos of, 113-138. 
rhinoceros, Oryctes. 
Rhizoctonus ampelinus, on Vitis, 66. 


Rhodesia, Saissetia subpatelliformis in, 
13; bionomics of Glossina morsitans 
in, 104, 113-132; new weevils from, 
2d hye os wt Oe 


Rhopalosiphum carduellinum, on Car- 
dwus in Transvaal, 67. 
Rhynchium cyanopterum, 149. 


Rhynchophorus ferrugineus, on date 
palms in India, 295. 


phoenicis, on date 
palms in Africa, 295. 


Rhyncosia memnonia, food-plant of 
thrips in Sudan, 98. 


richiardii, Taeniorhynchus. 
rima, Culex. 

ripariae, Aphis. 

ronaldi, Culex. 

rosae, Diaspis. 


rossi, Anopheles 
A. subpictus). 


rotunda, Forda. 
rubramarginata, Phara (Cadicera). 


99 


(Myzomyia) (see 


GENERAL 


rufa, Pangonia (see Corizoneura tapro- 
banes). 


rusci, Ceroplastes (Coccus). 


Russia, North, blood-sucking flies of, 
195-198. 


sabella, Arenipses. 
saccharina, Tomaspis. 
sacchariphaga, Proceras. 
sagittaria, Corizoneura. 
Saissetia, 2, 3. 
5 depressa (see S. nigra). 
sd hemisphaerica, in 8. Africa, 9. 


a kellyi, sp. n.,. on Acacia 
melanoxylon in Natal, 12. 


gs nigra, on Ficus in 8. Africa, 
10. 
4 nigrella (see S. nigra). 


i oculata, sp. N., on grape vine 
m: Natal, <b. 


9 (Lecanium) oleae, in Cape 
Colony, 10; on Phoenix, 
299. 

oF perseae, sp. 0., on Persea 
gratissima in Natal, 11. 

ed persimilis, on Combretum in 
Transvaal, 12. 

9 subpatelliformis, food-plant 


of, in Rhodesia, 13. 
salinarius, Culex. 
salisburiensis, Culex. 


Salt Water, mosquitos breeding in, 260, 
261. 


sanguinis, Hodgesia. 

Sarcopsylla penetrans, an introduced 
pest in Mauritius, 173. 

saussuret, Anoplodesmus. 


Schistocerca peregrina, attacking date 
palms in Sudan, 300. 


Schizoneura ampelorhiza, on Vitis, 56. 
schiffneri, Anopheles (Nyssorhynchus). 
schwetzi, Corizoneura. 

Scione alluaudi (see Bouvierella). 


Scirpus, Simulium breeding amongst, 
213, 232, 234, 235, 237. 


sculpturata, Calandra. 
scutellaris, Oremastogaster ; Stegomyia. 


Scymnus, predaceous on Parlatorta 
blanchardi, 297. 

Selenaspidus (Aspidiotus) articulatus, 
food-plants of, 299. 


INDEX. 431 


sepuleralis, Chrysops. 


sergenti, Anopheles culicifacies ; Phle- 
botomus. 


sericalum, Simulium. 
sericeum, Simulium. 
Setaria mauritiana, in §. E. Africa, 322. 


setifera, Lachnosterna ; Loncheaa (Car- 
polonchaea). 


Seychelles, Lonchaea in, 202 ; Tabanids 
from, 43—45. 


Shorea robusta, Calandra shoreae on, in 
India, 277. 


shoreae, Calandra. 
siberiensis, Theobaldia. 


Sideroxylon  dulcificum, 
breeding in, in Lagos, 257. 


signatipennis, Phlebotomus. 
Tecanium (see LI. hes- 


mosquitos 


signiferum, 
peridum). 

Silvanus surinamensis, infesting stored 
dates in Mesopotamia, 303. 


similliimus, Phlebotomus. 


simplex, Chionaspis ; Oligonychus (Para- 
tetranychus). 


simpsont, Stegomyia. 


simulans, Lecanium depressum (see 
L. hesperidum). 


in North Russia, 197; 
bionomics and classifica- 
tion of British species of, 
211-246; keys to British 
species of, 216-223; an- 

4 atomy of, 218, 220, 222; 

parasites of, 215. 


oh angustipes, the male of 
S. aureum, 217, 242, 243. 


pe angustitarsis, in Britain, 
DL7y) 218; 216, 221,. 222, 
223, 241, 243 ; previously 
described as S. latipes, 
239; synonomy of, 242, 
243. 


“3 arcticum, in British Colum- 
bia, 229. 


o argenteostriatum _ possibly 
spring form of S. argyrea- 
tum, 236. 


yy argyreatum, in Britain, 211, 
212,) 214.) 217, 218, 220, 
223, 232, 234, 237, 241. 


Simulium, 


™ argyreatum var. sericatum, 
in Britain, 214, 217, 218, 
235. 


432 


GENERAL INDEX. 


Simuliwm aureum, in Britain, 211-242; 


9? 


99 


2” 


99 


99 


Synonymy of, 243. 
aureosimile, in Africa, 244. 


austemi, a local race of 
S. venustum, 217, 218, 
232. 


botulibranchium, in South 
America, 238. 


bracteatum, Synonymy of, 
217, 243. 


columbaczense, doubt as io 
identity of, 242. 


damnosum, in Africa, 238. 

equinum, in Britain, 212- 
234, 236, 238-241. 

hirtipes, in Britain, 217, 
218, 220, 231, 245, 246: 
in North Russia, 198. 


latipes, in Britain, 218, 220, 
222, 228, 230, 241—245; 
in North Russia, 198. 


maculatum, 8. latipes not 
considered a synonym 
of, 241, 242. 


monticola, in Britain, 216, 
218; 220, 222) 222, 5226, 
228, 229. 


morsitans, in Britain, 217, 
218.) 221, 222, 227; 232; 
233. 


obreptans, 230. 


ornatum, in North Russia, 
198; bionomics of, in 
Britain,, 211—220, 223, 
224-231; parasites of, 
215. 


ornatum var. nitidifrons, 
Nz, jin, oribaim,. 218. 222. 
224, 226. 

piscicidium, in 
America, 225, 228. 

reptans, in Britain, 211-228, 
230, 234-239; in North 
Russia, 198. 


reptans var. galeratum, 2., 
probably identical with 
S. reptans, 218, 231. 


sericatum, in Britain, 211. 


North 


sericeum, 241; S. ornatum 
previously recorded as, 
226. 


subexcisum, 198; in Britain, 
217-223, 244, 245. 


Simulium subornatum, sp. n., in 
Britain, 211—227. 


= tredecimatum, sp. 0., in 
Britain, 215, 219, 221, 
222, 246. 

mi trifasciatum, a form of S. or- 
natum, 225. 

5 tuberosum, in Britain, 216, 
280, 234. 


variegatum, in Britain, 216— 
223, 228, 229, 230: 


Pe venefica, S. variegatum des- 
cribed under name of,228. 


venustum, in America, 238 ; 
in Britain, 213—223, 232, 
233.7 in {North Rissa, 


198. 
is yerburyi, sp. n., in Britain, 
219, 221, 222, 245. 
sinensis, Anopheles (Myzorhynchus) 


(see A. hyrcanus). 
Singapore, Lonchaea spp. in, 207, 209. 
singularis, Thriambeutes. 
sitiens, Culex. 
smithi, Lachnosterna (Phytalus). 


Sodium Fluoride, comparison of, with 
formaldehyde, 63. 


sorghi, Aphis. 


Solanum incanum, food-plant of thrips 
in Sudan, 98. 


PA wildeni, Coccids on, in 
Mauritius, 174. 


Solenopsis, an introduced pest in Mau- 
ritius, 173. 


sollicitans, Culex. 

solstitialis, Tabanus. 

soncht, Macrosyphum. 

Spain, Aphelinus mytilaspidis para- 
sitising Parlatoria blanchardi in, 297 ; 
Dociostaurus crucigerus in, 403. 


Sparganium, Simulium bre-ding 
amongst, 213, 224, 235, 241. 
spectabile, Asterolecanium. 


Spermatophora hornigu, on dates in 
Mesopotamia, 288. 


Sphaerococeus draperi 
cococcus marlatti). 


splendens, Mimomyia. 

splendida, Lonchaea (see L. aurea). 
squamipennis, Aéedomyia. 
squamosus, Anopheles. 


(see Phoeni- 


il 


GENERAL INDEX 


Stauronotus (see Dociostaurus). 


* annulipes (see Doctosta- 
urus crucigerus). 

i australis, 398. 

Fy genet (see Dociostaurus 
crucigerus). 

os hauensteint (see Dociosta- 
wrus crassiusculus). 

= kervillei (see Doctostaurus 
crassiusculus). 

PP tartarus, a geographical 
race of Docvostaurus 


crucigerus, 398, 403. 
Stegomyia africana, breeding places of, 
in Lagos, 256, 264, 265, 
270. 
" albopicta, 134 ; in Madagas- 
car and Réunion, 134. 


és apicoargentea, in Lagos, 269. 
me cartroni (see Aédes). 
= fasciata, 106; breeding 


places of, in Lagos, 248— 
270; in Mauritius, 134; 
in Palestine, Egypt and 


Syria, 392. 

we lamberti (see S. albopicta). 

Pe luteocephala, breeding places 
of, in Lagos, 251-270. 

fe metallica, breeding places 
of, in Accra, 258. 

5 pollinctor (see Ochlerotatus 
longipalpis). 

re scutellaris (see S. albopicta). 

ig simpsoni, breeding places 
of, in Lagos, 258. 

+ unilineata, breeding places 
of, in Gold Coast, 258, 
259. 


Stegonocops, breeding in salt water in 
Panama, 261. 


Stenobothrus (see Dociostaurus). 
Stenommatus fryi, in Mexico, 278. 


. musae, Sp. D., on banana 
in Hawaii, 277. 


Sterculia barteri, mosquitos breeding 
in, in Lagos, 257. 


= triphaca, in Glossina areas in 
S.E. Africa, 338. 


Stethomyia aitkeni (see Anopheles). 


Stomoxys, in S.E. Africa, 368; in 
North Russia, 197. 
A calcitrans, in North Russia, 


198. 


433- 


suavis, Buplex (Pangonia). 

subexcisum, Simulium. 

subfascia, Buplex (Pangonia). 
subornatum, Simulium. 
subpatelliformis, Saissetia (Lecanwm). 
subpictus, Anopheles (Myzomyia). 
subterranea, Conofilippia. 
subtruncatus, Elytroteimue. 


Sudan, bionomics and control of. 
Heliothrips indicus on cotton in, 95— 
100 ; Schistocerca peregrina on date 
palms in, 300, 301. 


Sugar-cane, bionomics and control of 
Lachnosterna (Phytalus) smithi on, in 
Mauritius, 159-169; Tomaspis sac- 
charina on, in Trinidad, 179. 


Sumatra, Lonchaea spp. in, 207, 209. 

swperpictus, Anopheles. 

surinamensis, Silvanus. 

sygmoides, Dacus. 

sylvestrii, Lecaniwm 
persimilis). 

Syria, mosquitos in, 390-395. 

Systates beiranus, sp. n., in East Africa,. 


(see Saissetia’ 


274. 

AS chirindensis, sp. n., on coffee: 
in Rhodesia, 275. 

= exaptus, 8p. n., On maize in 


Rhodesia, 273. 


Tabanids, classification of African, 
139-152; of Seychelles, 43; in 
North Russia, 195, 197, 198. 


Tabanus, in Portuguese East Africa, 
341, 374. 


oe aethiopicus (see Corizoneura). 
A! albipectus, in Seychelles, 45. 
di barbatus (see Osca). 


FP borealis, in North Russia,. 
198. 

< crocodilinus, in Nyasaland,. 
45. 


bs lapponicus, in North Russia, 
198. 


oe luridus, in North Russia,. 
198. 

a maculicornis, in North Rus-. 
sia, 196, 198. 

a nigricornis, in North Russia, 
198. 


ve nigrifacies, in North Russia,. 
196, 198. 


434 


Tabanus solstitialis, in North Russia, 


198. 

x tarandinus, in North Russia, 
196, 198. 

‘. tropicus, in North Russia, 
198. 

7 tropicus var. bisignatus, in 


North Russia, 198. 
tachardiaformis, Ceroplastes. 
Taeniorhynchus annetti, in Lagos, 268, 

270 


9 aureus, 135. 

5 aurites, in Lagos, 268, 
210. 

Me (Coquillettidia) flavus, 
in Madagascar, 135. 

9 fuscopennatus, 135. 

i” metallicus, in Lagos, 
269, 270. 

se richiardiui, recorded 
from Palestine, 395. 

5 uniformis (see Man- 


sonioides). . 
taeniorhynchus, Culex. 


tananariviensis, Anopheles (Cellia) (see 
A. squamosus). 

Tanyglossa aethiopica (see Corizoneura). 

Tanganyika ‘Territory, Selenaspidus 
(Aspidiotus) articulatus in, 299; 
Pangonia spp. in, 148, 151. 

Tanymecus agricola, sp. 1., on maize 

in Rhodesia, 272. 


- destructor, sp. 1., on maize 
and sweet potatoes in 
Rhodesia, 271. 


. inafectatus, 271. 
Tapinoma — erraticum — migerrimum, 
associated with Aphids in Persia, 
156. 
taprobanes, Corizoneura. 
tarandinus, Tabanus. 
tarsimaculatus, Anopheles. 
tartarus, Dociostaurus (Stauronotus). 
Tatera, in Glossina areas in S.E. Africa, 
338. 
Tephrosia emeroides, 
thrips in Sudan, 98. 
Terminalia catappa (African Almond), 
mosquitos breeding in, 
in Lagos, 257. 
" sericea, in relation to dis- 


tribution of Glossina in 
S.E. Africa, 349. 


food-plant of 


GENERAL INDEX 


terminaliae, Lecanium (see IL. hes- 


peridum). 
terticeps, Aegophagamyia. 
tessellatum, Lecanium. 


tessellatus, Anopheles (Nyssorhynchus) 
punctulatus. 


testudinata, Columnea (see Ceroplastes 
TUSCt). 


testudineus, Calypticus (see Ceroplastes 
TUSCL). 


testudiniformis, Columnea (see Cero- 
plastes rusct). 


testudo, Lecanium oleae (see Saissetia). 
thalassius, Culex. F 
Theobaldia annulata, in Palestine, 395 ; 
variety of, 393. 
é longiareolata, in Palestine, 
Egypt and Syria, 395. 


Es morsitans, larva of, in 
Palestine, 395. 

aw siberiensis, in North Russia, 
198. 


theobromae, Hemilecanium. 
Thriambeutes, 149. 


ry fuscus; sp. 2D, mm 
Bechuanaland, 151, 
152; 

ag singularis, 152. 


Thrips, on date palms in Trinidad, 295 ; 
on cotton in Sudan, 95—100. 


tigripes, Culex. 


Tipula oleracea, possible seasonal vari- 
ation of, 214. 


s»  paludosa, possibly a seasonal 
form of 7. oleracea, 214. 


»  parallela, establishment of, in 
Mauritius against Lachnos- 
terna (Phytalus) smithi, 159, 
163, 164, 168. 


titillans, Mansonia. 


Tomaspis saccharina, measures against, 
on sugar-cane in Trinidad, 179. 


Toxorhynchites brevipalpis, in Lagos, 
269 ; predaceous larva of, 281 (note). 


transvaalensis, Anopheles. 
tredecimatum, Simulium. 


Trees, mosquitos breeding in, in Lagos, 
255-259. 


Trichilia, in relation to distribution of 
Glossina in 8.E. Africa, 348, 349. 


trifasciatum, Simulium. 


GENERAL INDEX 


Trinidad, Heliothrips haemorrhoidalis 
on date palms in, 295 ; Lachnosterna 
spp. in, 192, 193; Lomaspis sac- 
charina in, 179. 


trinitatis, Lachnosterna. 
Triumfetta, in 8.E. Africa, 326. 
tropicus, Tabanus. 

Tsetse Flies (see Glossina). 
tuberosum, Simulium. 
turkhudi, Anopheles. 


Uapaca spp. influence of, on distribu- 
tion of Glossina in §.E. Africa, 320, 
326, 352, 365. 


Uganda, new Aphids from, 65—72. 
ulmi, Lepidosaphes. 

umbratipennis, Corizoneura. 
umbrosus, Anopheles (Myzorhynchus). 
unguiculata, Uranotaenia. 


uniformis, Mansonioides (Taeniorhyn- 
chus). 

unilineata, Stegomyia. 

United States of America, Parlatoria 
blanchardi and its parasites in, 288, 
297. 

univittatus, Culex. 

Uranotaenia annulata, breeding places 

of, in Lagos, 251-270. 
ea balfowri, in Lagos, 269, 
270. 


- bilineata var.  fraseri, 
breeding places of, in 
Lagos, 251-270. 


- mashonaensis, in Lagos, 
269. 

a neireti, sp. n., in Mada- 
gascar, 138. . 

mS ornata, breeding places 
of, in Sierra Leone, 
258. 

fn unguiculata, in Palestine 


and Egypt, 395. 


435 


vagus, Anopheles (Myzomyia). 
variegatum, Simuliwm. 

venefica, Simuleum. 

ventrale, Lecanium (see L. hesperidum). 
ventrilloni, Culex. 

venustum, Simulium. 


Vernonia podocoma, 
distribution of Glossina 
Africa, 322, 348. 


Vespa orientalis, destroying dates in 
Mesopotamia, 300. 


victrix, Parlatoria (see P. blanchardi). 


Villa Uoydi, parasite of Glossina 
morsitans in Rhodesia, 104. 


violae, Phorodon. 
virgata, Corizoneura. 


in relation to 
in: Be. 


viticola, Macrosiphum. 

Vitis, Aphids recorded on, 66. 
vitis, Aphis. 

vitisana, Peritymbia. 


Vitex eylesii, in relation to distribution 
of Glossina in S.E. Africa, 348. 


walkeri, Phlebotomus. 


Water Hyacinth (see WHichornia 
crassipes). 
avg Lettuce (see Pistia stratiotes). 


Websterella blanchardi (see Parlatoria). 


West Indies, new species of Lachno- 
sterna in, 189. 


wisteriae, Lecaniun. 
Ximenia americana, in §.E. Africa, 321. 
yerburyt, Simulium. 


zammittii, Acartomyia (Aédes). 

Zanzibar, Lonchaea (Oarpolonchaea) 
laevis in, 203; Icerya aegyptiaca on 
date palms in, 299. 

zizyphi, Inglisia ; Parlatoria. 

zonata, Pangonia. 

zonatus, Ceroplastes. 


7) 


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| icin a ie iain 
| me vy 
At 


INDEX TO NAMES 


437 


OF PERSONS. 


Adelung, 405. 

Aders, Dr. W. M., 45, 203. 

Alcock, Lt.-Col. A., 241. 

Aldrich, 214. 

Anderson, T. J., 204. 

Andrewes, H. L., 276. 

Armstrong, L., 204. 

Arrow, G. J., 189, 288, 292, 293, 294. 
Ashworth, Dr. J. H., 230, 237, 240. 
Atkinson, J. M., 281. 

Aurivillius, Prof. C., 277. 


Austen, Major E. E., 43, 103, 139, 195, 
tS7, S31. 


Azam, 404. 


Bacot, A., 256, 258. 
Bagnall, R. S., 95. 


Bagshawe, Dr. A. G., 317, 336, 362, 
367. 


Baker, Prof., 47. 

Baker, C. F., 207, 208, 209. 
Ballou, H. A., 191. 

Bates, H.W., 189, 192. 
Bedford, G. A. H., 233. 
Bedford, G. B., 67. 
Beeson, C. F. C., 276, 293. 
Bequaert, 362. 

Bezzi, Prof., 172, 199. 
Blacklock, 259. 

Blair, K. G., 225. 
Blanchard, 137. 

Bolivar, C., 397, 403, 406. 
Bolivar, I., 397, 403, 406. 
Bouet, 133, 135, 136, 137. 
Botha, L.-J., 7. 

Boyce, Sir R., 254. 

Brain, C. K., 1. 

Brenske, 189. 

Bréthes, 139. 


Britten, H., 212, 237. 
Brown, J. KE. M:, 276. 


| Buckler, W., 289, 301. 


Buddle, R., 198. 
Burkill, L.. H., 294, 301. 
Burmeister, 190. 
Busck, 73. 


Buxton, Capt. P. A., 154, 155, 156, 157, 
181, 287, 301, 305, 306, 310, 311. 


Byam, Lt.-Col. W., 53. 


Cajus, F., 208. 
Carment, Capt. A. G., 198. 


Carpenter, Dr. G. D. H., 145, 149, 151, 
341. 


Carr, Jc W., 227, 243. 
Carter, H. F., 259. 
Castellani, Dr., 226. 
Chalmers, Dr. A. J., 226. 
Champion, H. G., 277. 
Chand, 259. 


Christophers, Major 8S. R., 89, 90, 91, 
259, 389. 


Chittenden, F. H., 289, 301. 
Chubb, E. C., 318. 
Cockayne, Dr. E. A., 198. 


Cockerell, Prof. T. D. A., 15, 28,. 295, 
296, 297, 298, 301. 


Collins, Miss F., 233. 
Connal, Mrs., 250. 
Connal, Dr. A., 250. 
Cook, A. J., 301. 
Copeland, E. B., 294, 301. 
Coquerel, Dr., 45. 
Corbett, G. H., 95. 
Crawley, W. C., 154. 
Carty, Dr. 2. P:,. 73. 


d’Emmerez de Charmoy, D., 27, 133, 
134, 137; 159, 160, 171. 


438 


Dalziel, Dr. J. M., 247. 

De Meijere, Prof., 206, 207, 210, 240. 
Dowson, V. H. W., 303. 
Draper, W., 297, 301. 

Drége, 277. 

Dreyer, T. F., 40. 

Dudgeon, G. C., 289. 

Duke; Dr: Bot ane . 
Dupont, P. R.,. 45, 

Durrant, J. H., 288; 290, 30L. 
Dyar, H., 254, 261, 281. 


Edwards, FE’. W., 77, 105, 112, 133, 211, 
395. 


Elridge, E., 191. 

Enderlein, Dr. G., 244. 
Ensor, 336. 

Essig, E. O., 296, 299, 301. 
Evans, W., 225, 226. 

Evans, Capt. W. E., 293, 295. 
Eversmann, 403. 


Faure, J.C 24. 

Fernald, Mrs., 16. 

Ferreira, 317. 

Fiske, W. F., 117, 118. 

Forbes, R. H., 296, 297, 298, 299. 
Forbes, Dr. S. A., 74. 

Forbes, W. T. M., 186. 

Fracker, 8. B., 186, 290, 291, 301. 
Fraser, Major A. D., 195, 198. 


Fredericha, Dr. K., 21), 226; 2268; 230, 
242. 
Fryer, J. C. F., 43, 45. 


Fuller, C., 4, 15, 22, 24, 26, 27, 28, 31, 
65, 66,, G7. 


Garnett, H., 235, 243. 
Gardiner, Prof. J. S., 43, 45. 
Ghosh, C. C., 294, 301. 
Giese, A., 128. 

Giles, 89, 91. 

Godwin, Mrs., 11. 

Goodall, T. B., 141. 

Gough, Dr. L. H., 289, 302. 
Gowdey, C. C., 65, 71, 72. 


INDEX TO NAMES OF 


PERSONS. 


Graham, Dr. W. M., 256, 263, 266, 270, 
362. 


Green, E. E., 288, 299. 
Grimshaw, P. H., 231. 
Grinberg, 226. 

Gunn, D., 2s. 


Hagedorn, M., 302. 
Halford, F. M., 228. 
Hamilton, Major J. Stevenson, 339. 


| Hamm, A. H., 227, 233, 234, 238, 241, 


242, 243. 
Hampson, Sir G. F., 181, 182, 290, 302, 
Hearsey,, Dr.,. 337. 
Hensel, 214. 
Hesse, B. L., 225. 


| Ball, GG... 207. 


Hirst, S., 299, 300, 302. 

Hobson, J., 37. 

Holub, Dr., 128. 

Horvath, G., 243. 

Howard, Dr. L. O., 254, 261, 281. 
Hustache, Prof. A., 277. 


Ikonnikov, N., 397. 
Impey, Miss E., 30. 


Ingram, Dr. A., 74, 75, 105, 137, 258, 
259, 267, 305, 306, 311. 


Jack, R. W., 13,113, 152; 271) (272, 
273, 317, 318, 351, 353. 


Jackson, Miss A. C., 63. 
Jaillet, 136. 

Jardine, N. K., 209. 
Jenkinson, F., 246. 
Jenkyns, Lady, 243. 


Kandelaki, Dr., 307. 

Kelly, A. E., 6, 12, 17, 20, 22, 37, 39. 
Kemp, S. W., 276. 

Kertész, Dr., 204. 


King, H. H., 295, 296, 297, 300, 301, 
302. 


Kinghorn, Dr., 341. 
Kirby, W. F., 397. 
Knab, F. 73, 75, 254, 261, 281. 


INDEX TO NAMES OF PERSONS. 


La Baume, W., 402. 
Lamb, P. H., 204. 


Lamborn, Dr. W. A., 101, 114, 279, 
336, 358, 359, 364, 367, 376, 384. 


Lanne, 316, 317. 


Lawrence, Dr. W. T., 317, 329, 335, 
347, 351, 353, 354, 362, 363, 384. 


Lawrence, Mrs., 317, 346. 
Leatham, Capt. W. H., 305. 
Lefroy, Prof. H. M., 47, 63. 
Lesne, P., 192. 

Liebe, 226. 


Lloyd, Capt. Ll., 47, 130, 328, 336, 339, 
340, 341, 363, 364, 384. 


Lodge, Miss O. C., 47, 63. 
Longden, W. M., 318, 334. 
Lounsbury, C. P., 39, 296, 302. 
Ludlow, Miss, 89, 91. 

Lugard, Sir F., 152. 

Liitg, Dr: A., 211, 311: 


McConnell, Dr. R. E., 362. 


Macfie, Dr. J. W. 8., 74, 75, 105, 203, 
258, 267, 305, 306, 340. 


MacGregor, R. C., 207, 209. 
MacGregor, M. E., 283. 
Mackenzie, Dr. A., 123. 
Mackenna, J., 295, 302. 
Malloch, J. R., 211, 233, 245. 
Mally, C: W., 33, 63. 
Marzinovsky, 307. 

Marsden, E., 277. 


Marshall, Dr. G. A. K., 181, 271, 305, 
out, ‘aod. 


Maskell, W. M., 296, 302. 
Massey, R. E., 98. 
Maugham, R. C. F., 339, 340, 341, 358. 
Meinert, 226. 

Mennell, fF. P., 317, 325. 
Milne, D., 295, 302. 
Mochi, Dr. A., 202, 203. 
Moiser, Dr. B., 341. 
Monteiro, A. P., 317. 
Monteiro, J. J., 148. 
Morstatt, H., 299, 302. 


Moseley, M. E., 211, 215, 224, 228, 235, 
237, 238, 240, 241, 243. 


Munro, K., 20. 


439 


Newstead, Prof. R., 13, 19, 27, 212, 
237, 288, 297, 298, 299, 302, 305. 


Nesterov, P., 406. 

Neiret, Dr. 133, 136, 137. 
Nowell, W., 164. 

Nuttall, Prof. G. H. F., 389. 
Nurse, Lt-Col., 241. 


Old, Dr. J. EH. 8, 273. 

Olivia, L., 174. 

Orde-Browne, G. St. J., 204. 
Orner, A. J:, 352: 

Osmaston, A. E., 276. 
Otterson, G. D., 317, 337, 350. 


Peringuey, Dr. L., 41. 

Pettey, F. W., 35. 

Parrot, L., 307. 

Patterson, W. H., 204. 

Plotnikov, V., 397, 405. 

Pollard, Dr. J., 334. 

Pomeroy, A. W. J., 211, 233, 244. 


Popenoe, P. B., 292, 294, 296, 297, 300 
302. 


Portschinsky, Prof., 197. 


Ramme, Dr. W., 401. 
Reed, Miss B., 290. 
Rettie, Dr. J., 230, 240, 
Ritchie, A. H., 190. 
Romanoff, N. M., 302. 
Rothschild, Lord, 288. 
Roubaud, E., 364. 


Schwetz, Dr. J., 145, 146, 147. 
Scott, J. W., 317, 350. 

Séguy, E., 133, 135, 211, 235. 
Selous, Capt. F. C., 103, 128. 
Severin, G., 145. 

Sheppard, P. A., 274. 

Shipley, Sir A., 389. 

Shirecore, Dr., 346. 

Simpson, J. J., 358, 359. 

Sinton, Major J. A., 305, 306, 307, 309. 
Sloggett, Lieut.-Gen. Sir A. T., 143, 


| Smirnoy, D. A., 397. 


440 INDEX T NAMES OF PERSONS. 


Smyth, E. G., 189, 289, 302. 
Stebbing, E. P., 294, 302. 
Stephenson Hamilton, Major J., 339. 
Storey, G., 395. 

Strickland, C., 87, 91. 

Strickland, E. H., 215, 243. 

Stanton, Dr. A. Ta83,84, Ot. 
Staudinger, O., 186. 

Surcouf, J., 134, 137. 

Swellengrebel de Graaf, J. M. H., 77. 
Swellengrebel, N. H., 77, 92. 

Swezey, O. H., 276, 278. 
Swynnerton, C. F. M., 274, 275, 315. 


Tempany, H. A., 159, 191. 


Theobald, F. V., 65, 77, 153, 156, 260. 


Thompson, R. L., 272. 
Tomosvary, 242. 
Toumey, J. W., 302. 
Trabut, L., 298, 302. 


Urich,. F. W., 192, 193. 
Uvarov, Dr. B.?,, 306, 307, 397. 


Van der Goot, P., 155. 


Van der Merwe, C. P., 5, 9, 14, 19, 202. 


Van Heerden, 34. 


Van Leeuwen-Reijnvaan, Dr. J., 204. 


Van Leeuwen- Reijnvaan, Dr. W., 204. 


Ventrillon, Dr., 133, 134, 135, 136, 137, 
138. 


Verdat, 226. 
Vogler, 233, 238. 


Waldheim, F., 398. 
Walker, Dr. F. D., 311. 
Walton, C. L., 225, 228. 
Waterhouse, C. O., 397. 
Waterlot, 136, 137. 
Watermeyer, T. L., 16. 


Waterston, Capt. J., 238, 297, 305, 306, 
a07. 


Wenyon, Col. C. M., 307. 
Werner, F., 397. 
Whiteley, Dr. A. M., 49. 
Wilder, Dr:, 335. 


Williams, C. B., 179, 225, 228, 230, 241, 
295, 302. 


Wilsie, W. E., 297, 298, 302. 
Wimshurst, C. R., 303. 
Wilhelmi, Dr., 216. 
Willcocks, F. C., 261, 302. 


Yerbury, Lt-Col, J. W., 227, 243, 254. 


Zetek, J., 73. 
Zupitza, 362. 


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AUSTEN, Mas or H. E. (The Percy Sladen Trust Expedition to the 
Indian Ocean in 1908, and 1907-1909) Diptera : Tabanidae 


Brain, Dr. Cuas. K. The Chocstine ‘of South’ Africa—V. (Phares 
Ty bh eae Seat nN te rete RES UDI OS 


PAGE 


43. 


Luoyp, Capt. Lt. On the Rae for the Variation in. the © 


‘Effects of Formaldehyde as a Poison for House-flies - eh 


SWELLENGREBEL, Dr. N. H., & SWELLENGREBEL | DE Graar, J. M. H. 
List of the Anopheles of the Malay Archipelago with Special 


Reference to Adults and Larvae of New or POOR Ny 5 


Described Species or Varieties (illustrated) - ~- - = 
THEOBALD, Fre. VY. African Aphididae—Parr IV («lustrated) - 
Zurex, Jamus. Anopheles sail Among Water Lettuce —A 


New Habitat - = ii ied) Sok, eet ea ete 
MISOELLAN EOUS. 
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73 


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